P2050-004 exam Dumps Source : IBM Commerce Solutions Order Mgmt Technical Mastery Test v1
Test Code : P2050-004
Test cognomen : IBM Commerce Solutions Order Mgmt Technical Mastery Test v1
Vendor cognomen : IBM
: 30 true Questions
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ST. LOUIS--(business WIRE)--Perficient, Inc. (NASDAQ: PRFT) (“Perficient”), a leading digital transformation consulting company serving international 2000® and different gigantic commercial enterprise customers each and every through North the us, announced it has been named IBM’s 2019 Watson Commerce enterprise associate of the year. The IBM Excellence Award, announced right through IBM’s PartnerWorld at suppose 2019, recognizes Perficient’s ongoing increase and relationships with key shoppers, and notion management around the IBM Watson client rendezvous Commerce platform as an quintessential fraction for digital transformation.
“Our approach to commerce is concentrated on crafting a experience, connecting with purchasers, and supplying a seamless customer event across channels and each and every the way through the commercial enterprise, imperatives in nowadays’s buyer-pushed world,” mentioned Steve Gatto, countrywide sales director, Commerce solutions, Perficient Digital. “collectively, with their valued clientele, we’re remodeling agencies in a system that no longer best drives growth but strengthens their overall company, and they continually evolve their choices to retain customers at the excellent of their online game. We’re honored to live identified with the aid of IBM, and we’re anticipating sharing their innovative options during IBM Think 2019.”
Perficient Digital Takes Commerce solutions beyond Transactions to radically change the customer Lifecycle for a global varied brand
With branded manufacturers and distributors under pressure from the theatrical shift to online purchasing, a worldwide diverse company sought to digitally radically change its commerce business. In partnership with Perficient Digital, the two enterprises delivered optimized client income, updated product information (PIM), and streamlined the ordering technique through construction of a B2B portal. With the implementation of IBM’s Sterling Order administration device (OMS), and Perficient’s knowledge, the assorted brand is future-proofing its commerce to align with commerce trends and market opportunities.
furthermore, the business’s OMS will provide them more desirable flexibility in managing complicated order administration scenarios, stronger reliability so as processing and fulfilment, and a charge reduction in implementing across its enterprise. it's going to extra permit the organization to bring carrier enhancements to its purchasers, optimize its pricing, promoting and standard supply chain, boost earnings because of superior stock visibility, and slice back prices through better efficiencies so as visibility.
Perficient Digital Enhances the on-line consumer experience for a number one fabric Retailer
In a market that has historically trusted brick-and-mortar experiences, a number one cloth and craft retailer turned into challenged with extending the consumer adventure on-line. Perficient partnered with the commerce to result in force an IBM Watson Commerce retort that offered up to date visibility of its stock and greater tracking of its product amount, place, and availability. using IBM Order management, Perficient further more advantageous the solution via cloud migration that presents a single view of give and demand, orchestrates order success processes throughout buy on-line Pickup In save (BOPIS) and Ship-from-shop (SFS), and empowers commerce representatives to enhanced serve purchasers both in cognomen centers and in-save engagements.
“Perficient has been deploying IBM Commerce solutions for nearly 20 years, proposing conclusion-to-end digital commerce solutions that embody multiple channels, and convey seamless and efficient experiences throughout their entire business,” observed Sameer Peera, traditional supervisor, Perficient’s commerce practice. “With the contemporary information that HCL took over building of IBM WebSphere Portal, IBM internet content management and internet event manufacturing facility, their consumers proceed to engage us for aid with their digital commerce ideas. We’re completely blissful to live their go-to accomplice as they navigate the changing market landscape and deliver for his or her customers.”
Perficient learning in motion at IBM believe 2019
apart from its award-profitable commerce solution abilities, Perficient specialists are accessible throughout the IBM feel 2019 conference in sales space #320 to talk about its experience and capabilities across the IBM portfolio , chiefly cloud, cognitive, facts, analytics, DevOps, IoT, content management, BPM, connectivity, commerce, mobile, and customer engagement.
whereas IBM has introduced its plans to sell its commerce portfolio, the word of its acquisition of crimson Hat additionally signaled the criticality cloud construction and beginning play in a hit conclusion-to-conclusion digital transformations. As an IBM global Elite accomplice, one in every of most efficacious seven companions with that fame globally, and a purple Hat Premier partner, Perficient is smartly positioned to work with each companies via this transition. And, their consultants could live on hand each and every the way through IBM suppose to talk about the way to navigate the cloud market, share key client success reports, and provide strategic learning on the alternatives forward for valued clientele.
“technology is altering so unexpectedly, and firms necessity to preserve pace or mug disruption,” spoke of Hari Madamalla, vice chairman, emerging options, Perficient. “With learning and adventure in each and every features of the commerce journey, to leading cloud, internet hosting, managed capabilities and assist options, companies turn to Perficient as a go-to associate for his or her digital transformations.”
join a yoke of Perficient discipline depend consultants and their customers as they current each and every through six IBM suppose classes, together with:
As a Platinum IBM company partner, Perficient holds more than 30 awards throughout its 20-year partnership historical past. The company is an award-winning, certified utility cost Plus retort provider and one of the vital few companions to receive dozens of IBM professional stage software competency achievements.
For updates during the event and after, connect with Perficient experts online by way of viewing Perficient and Perficient Digital’s blogs, or comply with us on Twitter @Perficient and @PRFTDigital.
Perficient is the main digital transformation consulting enterprise serving global 2000® and commercial enterprise valued clientele during North the us. With unparalleled counsel technology, administration consulting, and inventive capabilities, Perficient and its Perficient Digital agency deliver imaginative and prescient, execution, and value with astonishing digital experience, enterprise optimization, and trade solutions. Their work allows consumers to enhance productiveness and competitiveness; develop and beget stronger relationships with shoppers, suppliers, and companions; and in the reduction of charges. Perficient's authorities serve clients from a network of offices throughout North the us and offshore areas in India and China. Traded on the Nasdaq international opt for Market, Perficient is a member of the Russell 2000 index and the S&P SmallCap 600 index. Perficient is an award-winning Adobe Premier partner, Platinum flush IBM enterprise accomplice, a Microsoft country wide provider company and Gold certified partner, an Oracle Platinum companion, an advanced Pivotal able partner, a Gold Salesforce Consulting partner, and a Sitecore Platinum associate. For more tips, seek counsel from www.perficient.com.
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probably the most statements contained during this information free up that aren't simply ragged statements focus on future expectations or state different forward-looking suggestions related to fiscal results and company outlook for 2018. these statements are realm to general and unknown dangers, uncertainties, and different components that could trigger the precise consequences to vary materially from these pondered by the statements. The forward-looking assistance is according to administration’s existing intent, belief, expectations, estimates, and projections concerning their company and their trade. bethink to live mindful that those statements handiest reflect their predictions. specific routine or results may additionally fluctuate considerably. essential factors that might understanding their precise outcomes to live materially distinctive from the ahead-searching statements embrace (however don't appear to live restricted to) those disclosed beneath the heading “possibility factors” in their annual document on configuration 10-k for the year ended December 31, 2017.
this might encompass IBM Watson Order management and Commerce for seamless digital engagement. Working with IBM enterprise colleague CEBS worldwide, IBM options will no longer simplest support power superior customer experiences and new ranges of comfort but convey efficiencies to the supply chain.
With a national footprint of 350 actual showrooms, an increasing manufacturer portfolio and changing client preferences, Metro footwear Ltd turned into facing challenges in managing orders coming from dissimilar on-line structures.
past it had unreliable software that brought about want of visibility of real-time information of income, inventory region and returns together with inventory management challenges. Metro footwear Ltd vital to enrich online presence for some of their widely wide-spread interior brands which were getting low visibility impacting ordinary income."expertise is redefining client rendezvous and will live the famous thing differentiator for retail manufacturers of the long run. We’re excited to collaborate with IBM and CEBS to embark on their digital transformation event,” said Alisha Malik, vp, Digital, Metro shoes.
“With IBM’s competencies in the omni-channel commerce and retail area, they are assured that these adjustments will not handiest aid speed up the execution of their method, however furthermore give us an locality over competition. At Metro shoes, they strongly believe that the new solution will increase the general user experience, thereby increasing revisits, traffic and loyalty,” brought Malik.
With IBM, Metro shoes Ltd can benefit new levels of client perception, which may furthermore live used to customize the online adventure for each of the web site. enterprise will capable of demonstrate off each and every of its manufacturers and advocate specific items in accordance with insights shared through customers on a single platform.
This customized experience will embrace new and effortless fulfillment alternate options corresponding to purchase on-line, select up in save, reserve in save and simple returns. as a result of these new capabilities, Metro shoes should live able to raise every visitor’s adventure on the web site by way of enabling commerce practitioners with cognitive tools which aid them bring omni-channel experiences that hold interaction valued clientele and power earnings.
With IBM’s know-how capabilities and CEBS learning with market integration, Metro footwear as a company/seller will furthermore live in a position to integrate with more than 14 e-marketplaces like Amazon, Flipkart and other main portals with a centralized procedure and inventory engine to permit Metro to scale as much as the wants of a becoming marketplace business. further, IBM Cloud will support elevate the skill to configure cumbersome workloads and thereby convey performance required for height usage each and every the way through the searching season.
Nishant Kalra, company unit chief – IBM Watson consumer rendezvous - India/South Asia delivered, “IBM is at the forefront of helping consumers embody more recent how you can work and digitally remodeling the way they interact with their conclusion valued clientele. we're blissful to live fraction of Metro shoes’ digital transformation adventure via delivering sophisticated digital commerce adventure, leveraging the stores via merging them with online, and at last driving company advocacy. IBM in affiliation with CEBS will permit profound innovation, quicker-go-to-market and streamline processes for scalability.”
The IBM platform will create a bridge between its online and offline company which the retailer up to now lacked. With the new built-in single view, Metro footwear sooner or later will live capable of employ insights won from the digital realm to design special providing for clients as they stroll into any of their outlets. in consequence, they can account what purchasers want, ensure availability when and where they want it and even examine cross promoting and upselling throughout their a number of brands.
For Metro footwear, IBM Watson Order management and Commerce solutions can pave approach for IBM’s cognitive technologies to deliver insights that befriend them provide purchasers with personalised recommendations and an more desirable consumer journey –from click on to beginning.
“With over 15 years of event in developing e-business tools, CEBS has been a relied on options issuer and associate for businesses across the globe,” eminent Satish Swaroop, President, CEBS global. Their beneficial and supple application options paired with IBM’s profound technology talents will provide Metro shoes a real-time, centralized gadget for customer administration.”
aspect Roberts, WA and Vancouver, BC - February 14, 2019 (Investorideas.com Newswire) Investorideas.com (www.investorideas.com), a world investor information source overlaying artificial Intelligence (AI) brings you today's edition of The AI Eye - gazing inventory news, deal tracker and advancements in artificial intelligence.
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The #AI Eye: IBM ( $IBM) to invest $50 Million in analysis With sanatorium and medical middle and Qualcomm ($QCOM)
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modern day Column - The AI Eye - staring at stock information, deal tracker and developments in artificial intelligenceworld AI Chipset Market to strategy $13.4 Billion USD by using 2023
intellect Commerce has posted new research by means of In For boom, indicating that the world AI chipset market will system $13.four billion USD by means of 2023, with the united states producing over $three.5 billion. An excerpt from the document summary reads:
The AI chipset marketplace is poised to seriously change the entire embedded materiel ecosystem with a big number of AI capabilities comparable to profound laptop getting to know, photograph detection, and a lot of others. this may furthermore live transformational for current faultfinding enterprise functions corresponding to id management, authentication, and cybersecurity.
Multi-processor AI chipsets live trained from the atmosphere, users, and machines to uncover hidden pattern amongst facts, predict actionable perception, and fulfill movements in keeping with specific situations. AI chipsets will turn into an essential component of each AI utility/systems in addition to crucial assist of any records-intensive operation as they enormously enrich processing for a number of functions in addition to enhance universal computing performance.IBM to beget investments $50 Million in research With hospital and medical seat and Qualcomm, Accenture and Kellogg team to develop VR Merchandising solution stocks discussed: (NYSE:IBM) (NasdaqGS:QCOM) (NYSE:ACN) (NYSE:ok)
IBM, (NYSE:IBM) through Watson fitness, plans to beget investments $50 million in analysis collaborations with Brigham and girls's medical institution, a instructing health seat of Harvard scientific college, and Vanderbilt school clinical middle - to enhance the science of artificial intelligence (AI) and its software to main public fitness considerations. Work with the two associations will live cognizant of constructing options to fitness issues gold standard consummate for AI. Kyu Rhee, M.D., M.P.P., vp and chief fitness officer at IBM Watson fitness, commented:
"building on the MIT-IBM Watson Lab announced last yr, this collaboration will embrace contributions from IBM Watson fitness's lengthy-standing commitment to scientific analysis and their credit that working along with the world's main associations is the fastest course to strengthen, strengthen, and endure in mind purposeful solutions that transparent up probably the most world's biggest fitness challenges."
QUALCOMM incorporated (NasdaqGS:QCOM) subsidiary Qualcomm technologies, Accenture (NYSE:ACN) and Kellogg enterprise (NYSE:okay) are collaborating to better and pilot a digital truth merchandise answer. The VR merchandising solution makes employ of a Qualcomm VR reference design headset, powered with the aid of Qualcomm Snapdragon 845 mobile VR Platform and is developed by way of the Accenture extended reality (XR) practice. checking out turned into finished in collaboration with Kellogg across the launch of their Pop muffins Bites product. Raffaella digital camera, world head, Innovation & Market approach, Accenture extended fact, defined the tech:
"Our VR merchandising solution has the advantage to seriously change product placement by means of inspecting purchaser purchasing habits in a holistic approach. by means of combining the vigor of VR with eye-monitoring and analytics capabilities, it enables massive new insights to live captured whereas consumers store through monitoring the station and the way they evaluate each and every items across a whole shelf or aisle. finally, this permits product placement choices to live made that may positively hold an impact on total brand sales, versus simplest single product revenue."
Sam Mowers, Investorideas.com
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The example desktop management system should provide a "push" technology that allows administrators to deploy software to multiple PCs simultaneously from a centralized administrative console, without requiring quit user intervention or a technician to visit the desktop. Deployment tasks can live executed immediately or scheduled for off-hours in order to minimize impact on quit user productivity or network bandwidth.
The example desktop management should live an open and scalable system that supports a scope of server platforms, such as Solaris, HP-UX, NT, and both new and legacy Microsoft client platforms (DOS, Windows 3.x, Windows 95, Windows 98 and NT 4.0). The system should live standards-based, with support for standard protocols, including IP, DHCP and BOOTP and standard Wired for Management (WfM)-enabled PC platforms (DMI 2.0, Remote Wake Up and PXE). The desktop management system should furthermore support legacy PCs via boot PROMs or boot floppies for standard NICs from Intel, 3Com, SMC and others.
Essential to the equation should furthermore live a series of open, programmable interfaces that allow customers and partners to extend and customize the system. The system should live carefully designed to provide scalability across big numbers of clients and servers, including the competence to group PCs and software packages into deployment groups and the competence to intelligently manage network bandwidth.
Windows 2000 promises to address many of these limitations but will not live deployed in most production environments until 2001, according to industry analysts, such as the GartnerGroup; moreover, in order to buy advantage of these new desktop capabilities, organizations must migrate to an exclusive, all-Windows 2000 environment on both clients and servers, which may live unrealistic for many corporations, the preponderance of non-NT desktops.
The example desktop management system should configure operating systems, applications and desktop parameters on an ongoing basis. These operations should live executed simultaneously on multiple PCs from central administrative consoles, and should deliver three faultfinding capabilities: pre-OS installation, remote support and no quit user intervention. These three powerful capabilities result in enterprise desktop management nirvana: lower PC total cost of ownership (TCO).
As computing environments prance toward increasingly distributed and heterogeneous environments, many IT organizations are now implementing centralized management systems for managing network resources such as routers and printers, application and database servers (e.g., SAP, Oracle, Lotus Domino), and desktop PCs.
The driving force behind these implementations is the realization that centralized management systems are required to cost-effectively manage the complicated and mission-critical nature of networked systems. For most IT organizations, centralized management systems are the only way of approaching the same flush of reliability, availability and control as has been available with mainframe environments of the past.
Centralized desktop management tools are seen as a key requirement for reducing the TCO associated with desktop support and the rapid growth of desktops in enterprise environments, and as a key enabler for delivering a higher quality of IT service to end-user organizations.
In addition, most IT organizations now contemplate PC desktops as a mission-critical corporate resource that should live managed as fraction of an overall networked environment – embodying the philosophy "the network is the computer" – rather than treated as a series of isolated standalone resources to live managed on an individual basis.
Tactical requirements for desktop management typically arise in connection with urgent short-term projects such as desktop OS migrations (e.g., from Windows 3.1 or OS/2 to Windows 95/98 or NT), Y2K desktop remediation projects, large-scale deployments of new and more powerful PC hardware to support commerce unit requirements (Web access, e-commerce, multi-media, etc.), or deployment of new and complicated applications, such as Lotus Notes or Netscape Communicator.
A successful desktop management system should provide three key technology differentiators versus conventional electronic software distribution systems: pre-OS technology, native installation engine and continuous configuration.
The competence to install and configure operating systems on PCs that are new or are unable to boot due to corruption or misconfiguration is called pre-OS capability. Pre-OS technology enables the desktop management system to install operating systems on a PC regardless of its state (e.g., corrupted hard disk, won’t boot, virgin hard drive, etc.). If a desktop management system cannot fulfill these functions, then its value is tremendously reduced, as the (re)installation represents a major task of IT support staffs.
Pre-OS technology takes control of the PC even in the absence of a working operating system, and automates the installation and configuration of operating systems on new PCs out of the box. It furthermore acts in a befriend desk setting for PCs that are unable to boot due to misconfiguration or corruption – without requiring a technician to visit the desktop or any end-user interaction.
The example desktop management system should install applications by running the vendor-supplied native installation program (setup.exe) on the target client. Its desktop agent should click through the installation wizard using the installation options specified by the administrator before launching the installation task. This allows each installation to live easily customized on a per-user or group-wide basis via a point-and-click administrative interface. No editing of script or batch files is required. In addition, this approach provides a high flush of reliability because it leverages the vendor-supplied installation procedure that adapts in real-time to the hardware and software configuration of the target system.
The example desktop management system should manage PC configurations across the entire PC lifecycle, not just during the initial application installation. It should live able to deploy action packages to add a new printer or change printer settings, change the IP address or login password of a PC, flee an anti-virus or inventory scan, or execute a BIOS glitter as fraction of a Y2K remediation effort.
It is furthermore helpful for a desktop management system to maintain a unique client configuration database that stores a history of each and every software packages that hold been installed, as well as the configuration parameters that were used during installation. This database can live used to rebuild the desktop to its previous configuration at any time, in a completely unattended manner.
Intel WfM Initiative
The Intel WfM initiative is intended to significantly enhance manageability and reduce TCO for desktop PCs. According to Intel, approximately 14 million WfM-enabled PCs hold shipped since the quit of 1998.
WfM V2 will proffer enhanced manageability for mobile PCs, enhanced security via encryption and authentication, and support for new hardware/software asset management standards such as CIM (Common Information Model) and WBEM (Web-Based Enterprise Management). WfM V2 is currently in beta with PC manufacturers and is expected to live available in mid-1999.
In addition, 100 percent of the commerce PCs offered from vendors, such as Dell, Compaq, IBM and HP are currently shipping with WfM capabilities. The example desktop management solution should fully support the WfM V1.1 specification, which consists of three components:
Remote Wake Up (RWU): Allows IT organizations to execute administrative tasks remotely during off-hours to preserve network bandwidth and user productivity.
The PC client is automatically "awakened" under centralized control of the desktop management system, and directed to install and configure operating systems and applications.
DMI 2.0 (Desktop Management Interface): Developed by the Desktop Management Task force (DMTF), DMI 2.0 allows befriend Desk personnel to scan the hardware and software properties of remote PCs in real-time to aid in troubleshooting.
Knowledge of population connectivity is necessary for efficacious management in marine environments (Mitarai, Siegel & Winters, 2008; Botsford et al., 2009; Toonen et al., 2011). For many species of marine invertebrate and reef fish, dispersal is mostly limited to the pelagic larval life stage. Therefore, an understanding of larval dispersal patterns is faultfinding for studying population dynamics, connectivity, and conservation in the marine environment (Jones, Srinivasan & Almany, 2007; Lipcius et al., 2008; Gaines et al., 2010; Toonen et al., 2011). Many coastal and reef species hold a bi-phasic life history in which adults array limited geographic scope and high site fidelity, while larvae are pelagic and highly mobile (Thorson, 1950; Scheltema, 1971; Strathmann, 1993; Marshall et al., 2012). This life history strategy is not only common to sessile invertebrates such as corals or limpets; many reef fish species hold been shown to hold a home scope of <1 km as adults (Meyer et al., 2000; Meyer, Papastamatiou & Clark, 2010). Depending on species, the mobile planktonic stage can last from hours to months and has the potential to transport larvae up to hundreds of kilometers away from a site of origin (Scheltema, 1971; Richmond, 1987; Shanks, 2009). learning of larval dispersal patterns can live used to inform efficacious management, such as marine spatial management strategies that sustain source populations of breeding individuals capable of dispersing offspring to other areas.
Both biological and physical factors impact larval dispersal, although the relative import of these factors is likely variable among species and sites and remains debated (Levin, 2006; Paris, Chérubin & Cowen, 2007; Cowen & Sponaugle, 2009; White et al., 2010). In situ data on pelagic larvae are sparse; marine organisms at this life stage are difficult to capture and identify, and are typically institute in low densities across big areas of the open ocean (Clarke, 1991; Wren & Kobayashi, 2016). A variety of genetic and chemistry techniques hold therefore been developed to estimate larval connectivity (Gillanders, 2005; Leis, Siebeck & Dixson, 2011; Toonen et al., 2011; Johnson et al., 2018). Computer models informed by realm and laboratory data hold furthermore become a valuable tool for estimating larval dispersal and population connectivity (Paris, Chérubin & Cowen, 2007; Botsford et al., 2009; Sponaugle et al., 2012; Kough, Paris & Butler IV, 2013; Wood et al., 2014). Individual-based models, or IBMs, can incorporate both biological and physical factors known to influence larval movement. Pelagic larval duration (PLD), for example, is the amount of time a larva spends in the water column before settlement and can vary widely among or even within species ( Toonen & Pawlik, 2001). PLD affects how far an individual can live successfully transported by ocean currents, and so is expected to directly impress connectivity patterns (Siegel et al., 2003; Shanks, 2009; Dawson et al., 2014). In addition to PLD, adult reproductive strategy and timing (Carson et al., 2010; Portnoy et al., 2013), fecundity (Castorani et al., 2017), larval mortality (Vikebøet al., 2007), and larval developmental, morphological, and behavioral characteristics (Paris, Chérubin & Cowen, 2007) may each and every play a role in shaping connectivity patterns. Physical factors such as temperature, bathymetry, and current direction can furthermore substantially influence connectivity (Cowen & Sponaugle, 2009). In this study, they incorporated both biotic and abiotic components in an IBM coupled with an oceanographic model to predict fine-scale patterns of larval exchange around the island of Moloka‘i in the Hawaiian archipelago.
The main Hawaiian Islands are located in the middle of the North Pacific Subtropical Gyre, and are bordered by the North Hawaiian Ridge current along the northern coasts of the islands and the Hawaii Lee Current along the southern coasts, both of which flee east to west and are driven by the prevalent easterly trade winds (Lumpkin, 1998; Friedlander et al., 2005). The Hawai‘i Lee Countercurrent, which runs along the southern perimeter of the chain, flows west to east (Lumpkin, 1998). The pattern of mesoscale eddies around the islands is complicated and varies seasonally (Friedlander et al., 2005; Vaz et al., 2013).
Hawaiian marine communities mug unprecedented pressures, including coastal development, overexploitation, disease, and increasing temperature and acidification due to climate change (Smith, 1993; Lowe, 1995; Coles & Brown, 2003; Friedlander et al., 2003; Friedlander et al., 2005; Aeby, 2006). Declines in Hawaiian marine resources quarrel for implementation of a more holistic approach than traditional single-species maximum sustainable yield techniques, which hold proven ineffective (Goodyear, 1996; Hilborn, 2011). There is a general movement toward the employ of ecosystem-based management, which requires learning of ecosystem structure and connectivity patterns to establish and manage marine spatial planning areas (Slocombe, 1993; Browman et al., 2004; Pikitch et al., 2004; Arkema, Abramson & Dewsbury, 2006). Kalaupapa National Historical Park is a federal marine protected locality (MPA) located on the north shore of Moloka‘i, an island in the Maui Nui complicated of the Hawaiian archipelago, that includes submerged lands and waters up to 1 4 mile offshore (NOAA, 2009). At least five IUCN red-listed coral species hold been identified within this area (Kenyon, Maragos & Fenner, 2011), and in 2010 the Park showed the greatest fish biomass and species diversity out of four Hawaiian National Parks surveyed (Beets, Brown & Friedlander, 2010). One of the major benefits expected of MPAs is that the protected waters within the locality provide a source of larval spillover to other sites on the island, seeding these areas for commercial, recreational, and subsistence fishing (McClanahan & Mangi, 2000; Halpern & Warner, 2003; Lester et al., 2009).
In this study, they used a Lagrangian particle-tracking IBM (Wong-Ala et al., 2018) to simulate larval dispersal around Moloka‘i and to estimate the larval exchange among sites at the scale of an individual island. They hold parameterized their model with biological data for eleven species covering a breadth of Hawaiian reef species life histories (e.g., habitat preferences, larval behaviors, and pelagic larval durations, Table 1), and of interest to both the local community and resource managers. Their goals were to examine patterns of species-specific connectivity, characterize the location and relative magnitude of connections around Moloka‘i, recount sites of potential management relevance, and address the question of whether Kalaupapa National Historical Park provides larval spillover for adjacent sites on Moloka‘i, or connections to the adjacent islands of Hawai‘i, Maui, O‘ahu, Lana‘i, and Kaho‘olawe.Table 1:
Target taxa selected for the study, based on cultural, ecological, and/or economic importance.PLD = pelagic larval duration. Short dispersers (3–25 day minimum PLD) in white, medium dispersers (30–50 day minimum PLD) in light gray, and long dispersers (140–270 day minimum PLD) in unlit gray. Spawn season and timing from traditional ecological learning shared by cultural practitioners on the island. Asterisk indicates that congener-level data was used. Commonname Scientific name Spawn type # of larvae spawned Spawningday of year Spawning hour of day Spawning moon phase Larval depth (m) PLD (days) Habitat ’Opihi/ Limpet Cellana spp. Broadcast1 861,300 1–60 & 121–181 – New 0–5 3–181,2 Intertidal1 Ko’a/ Cauliflower coral Pocillopora meandrina Broadcast3 1,671,840 91–151 07:15–08:00 Full 0–54 5–90*5 Reef He’e/ Octopus Octopus cyanea Benthic6 1,392,096 1–360 – – 50–100 216 Reef, rubble7 Moi/ Pacific threadfin Polydactylus sexfilis Broadcast 1,004,640 152–243 – – 50–1008 259 Sand10 Uhu uliuli/ Spectacled parrotfish Chlorurus perspicillatus Broadcast 1,404,792 152–212 – – 0–120*11 30*12 Reef10 Uhu palukaluka/ Reddlip parrotfish Scarus rubroviolaceus Broadcast 1,404,792 152–212 – – 0–120*11 30*12 Rock, reef10 Kumu/ Whitesaddle Goatfish Parupeneus porphyreus Broadcast 1,071,252 32–90 – – 0–50*11 41–56*12 Sand, rock, reef10 Kole/ Spotted surgeonfish Ctenochaetus strigosus Broadcast 1,177,200 60–120 – – 50–10011 50*12 Rock, reef, rubble10 ‘Ōmilu/ Bluefin trevally Caranx melampygus Broadcast 1,310,616 121–243 – – 0–80*11 140*13,14 Sand, reef10 Ulua/ Giant trevally Caranx ignoblis Broadcast 1,151,040 152–243 – Full 0–80*11 14013,14 Sand, rock, reef10 Ula/ Spiny lobster Panulirus spp. Benthic15 1,573,248 152–243 – – 50–10016 27017 Rock, pavement16 Methods Circulation model
We selected the hydrodynamic model MITgcm, which is designed for the study of dynamical processes in the ocean on a horizontal scale. This model solves incompressible Navier–Stokes equations to recount the motion of viscous fluid on a sphere, discretized using a finite-volume technique (Marshall et al., 1997). The one-km resolution MITgcm domain for this study extends from 198.2°E to 206°E and from 17°N to 22.2°N, an locality that includes the islands of Moloka‘i, Maui, Lana‘i, Kaho‘olawe, O‘ahu, and Hawai‘i. While Ni‘ihau and southern Kaua’i furthermore descend within the domain, they discarded connectivity to these islands because they lie within the 0.5° restrict zone of the current model. restrict conditions are enforced over 20 grid points on each and every sides of the model domain. Vertically, the model is divided into 50 layers that increase in thickness with depth, from five m at the surface (0.0–5.0 m) to 510 m at the ground (4,470 –4,980 m). Model variables were initialized using the output of a Hybrid Coordinate Ocean Model (HYCOM) at a horizontal resolution of 0.04° (∼four km) configured for the main Hawaiian Islands, using the general Bathymetric Chart of the Oceans database (GEBCO, 1/60°) (Jia et al., 2011).
The simulation runs from March 31st, 2011 to July 30th, 2013 with a temporal resolution of 24 h and shows seasonal eddies as well as persistent mesoscale features (Fig. S1). They accomplish not embrace tides in the model due to temporal resolution. Their model age represents a neutral ocean state; no El Niño or La Niña events occurred during this time period. To ground-truth the circulation model, they compared surface current output to real-time trajectories of surface drifters from the GDP Drifter Data Assembly seat (Fig. S2) (Elipot et al., 2016), as well as other current models of the locality (Wren et al., 2016; Storlazzi et al., 2017).Biological model
To simulate larval dispersal, they used a modified version of the Wong-Ala et al. (2018) IBM, a 3D Lagrangian particle-tracking model written in the R programming language (R Core Team, 2017). The model takes the aforementioned MITgcm current products as input, as well as shoreline shapefiles extracted from the plenary resolution NOAA Global Self-consistent Hierarchical High-resolution Geography database, v2.3.0 (Wessel & Smith, 1996). Their model included 65 land masses within the geographic domain, the largest being the island of Hawai‘i and the smallest being Pu‘uki‘i Island, a 1.5-acre islet off the eastern coast of Maui. To model depth, they used the one arc-minute-resolution ETOPO1 bathymetry, extracted using the R package ‘marmap’ (Amante & Eakins, 2009; Pante & Simon-Bouhet, 2013).
Each species was simulated with a separate model run. Larvae were modeled from spawning to settlement and were transported at each timestep (t = 2 h) by advection-diffusion transport. This transport consisted of (1) advective displacement caused by water flow, consisting of east (u) and north (v) velocities read from daily MITgcm files, and (2) additional random-walk displacement, using a diffusion constant of 0.2 m2/s−1 (Lowe et al., 2009). Vertical velocities (w) were not implemented by the model; details of vertical larval movement are described below. Advection was interpolated between data points at each timestep using an Eulerian 2D barycentric interpolation method. They chose this implementation over a more computationally intensive interpolation system (i.e., fourth-order Runge–Kutta) because they did not celebrate a dissimilarity at this timestep length. Biological processes modeled embrace PLD, reproduction timing and location, mortality, and ontogenetic changes in vertical distribution; these qualities were parameterized via species-specific data obtained from previous studies and from the local fishing and management community (Table 1).
Larvae were released from habitat-specific spawning sites and were considered settled if they fell within a roughly one-km contour around reef or intertidal habitat at the quit of their pelagic larval duration. Distance from habitat was used rather than water depth because Penguin Bank, a relatively shallow bank to the southwest of Moloka‘i, does not picture suitable habitat for reef-associated species. PLD for each larva was a randomly assigned value between the minimum and maximum PLD for that species, and larvae were removed from the model if they had reached their PLD and were not within a settlement zone. No data on pre-competency age were available for their study species, so this parameter was not included. Mortality rates were calculated as larval half-lives; e.g., one-half of each and every larvae were assumed to hold survived at one-half of the maximum PLD for that species (following Holstein, Paris & Mumby, 2014). Since their focus was on potential connectivity pathways, reproductive rates were calibrated to allow for saturation of feasible settlement sites, equating from ∼900,000 to ∼1,7000,000 larvae released depending on species. Fecundity was therefore derived not from biological data, but from computational minimums.
Development, and resulting ontogenetic changes in behavior, is specific to the life history of each species. Broadcast-spawning species with weakly-swimming larvae (P. meandrina and Cellana spp., Table 1) were transported as passive particles randomly distributed between 0–5 m depth (Storlazzi, Brown & Field, 2006). Previous studies hold demonstrated that fish larvae hold a high degree of control over their vertical position in the water column (Irisson et al., 2010; Huebert, Cowen & Sponaugle, 2011). Therefore, they modeled broadcast-spawning fish species with a 24-hour passive buoyant facet to simulate eggs pre-hatch, followed by a pelagic larval facet with a species-specific depth distribution. For C. ignoblis, C. melampygus, P. porphyreus, C. perspicillatus, and S. rubroviolaceus, they used genus-level depth distributions (Fig. S3) obtained from the 1996 NOAA ichthyoplankton vertical distributions data report (Boehlert & Mundy, 1996). P. sexfilis and C. strigosus larvae were randomly distributed between 50–100 m (Boehlert, Watson & Sun, 1992). Benthic brooding species (O. cyanea and Panulirus spp.) accomplish not hold a passive buoyant phase, and thus were released as larvae randomly distributed between 50–100 m. At each time step, a larva’s depth was checked against bathymetry, and was assigned to the nearest available layer if the species-specific depth was not available at these coordinates.
For data-poor species, they used congener-level estimates for PLD (see Table 1). For example, there is no estimate of larval duration for Caranx species, but in Hawai‘i peak spawning occurs in May–July and peak recruitment in August–December (Sudekum, 1984; Longenecker, Langston & Barrett, 2008). In consultation with resource managers and community members, a PLD of 140 days was chosen pending future data that indicates a more accurate pelagic period.Habitat selection
Spawning sites were generated using data from published literature and modified after input from native Hawaiian cultural practitioners and the Moloka‘i fishing community (Fig. 1). Species-specific habitat suitability was inferred from the 2013–2016 Marine Biogeographic Assessment of the Main Hawaiian Islands (Costa & Kendall, 2016). They designated coral habitat as areas with 5–90% coral cover, or ≥1 site-specific coral species richness, for a total of 127 spawning sites on Moloka‘i. Habitat for reef invertebrates followed coral habitat, with additional sites added after community feedback for a total of 136 sites. Areas with a predicted reef fish biomass of 58–1,288 g/m2 were designated as reef fish habitat (Stamoulis et al., 2016), for a total of 109 spawning sites. Sand habitat was designated as 90–100% uncolonized for a total of 115 sites. Intertidal habitat was designated as any rocky shoreline locality not covered by sand or mud, for a total of 87 sites. Number of adults was assumed equal at each and every sites. For regional analysis, they pooled sites into groups of two to 11 sites based on benthic habitat and surrounding geography (Fig. 1A). Adjacent sites were grouped if they shared the same benthic habitat classification and prevalent wave direction, and/or were fraction of the same reef tract.Figure 1: Spawning sites used in the model by species. (A) C. perspicillatus, S. rubroviolaceus, P. porphyreus, C. strigosus, C. ignoblis, and C. melampygus, n = 109; (B) P. meandrina, n = 129;(C) O. cyanea and Panulirus spp., n = 136; (D) P. sexfilis, n = 115; and (E) Cellana spp., n = 87. Region names are displayed over associated spawning sites for fish species in (A). Regions are made up of two to 11 sites, grouped based on coastal geography and surrounding benthic habitat, and are designated in (A) by adjacent colored dots. Kalaupapa National Historical Park is highlighted in light green in (A). Source–sink dynamics and local retention
Dispersal distance was measured via the distm duty in the R package ‘geosphere’, which calculates distance between geographical points via the Haversine formula (Hijmans, 2016). This distance, measured between spawn and settlement locations, was used to compute dispersal kernels to examine and compare species-specific distributions. They furthermore measured local retention, or the percentage of successful settlers from a site that were retained at that site (i.e., settlers at site A that originated from site A/total successful settlers that originated from site A). To estimate the role of specific sites around Moloka‘i, they furthermore calculated a source–sink index for each species (Holstein, Paris & Mumby, 2014; Wren et al., 2016). This index defines sites as either a source, in which a site’s successful export to other sites is greater than its import, or a sink, in which import from other sites is greater than successful export. It is calculated by dividing the dissimilarity between number of successfully exported and imported larvae by the sum of each and every successfully exported and imported larvae. A value <0 indicates that a site acts as a net sink, while a value >0 indicates that a site acts as a net source. While they measured successful dispersal to adjacent islands, they did not spawn larvae from them, and therefore these islands picture exogenous sinks. For this reason, settlement to other islands was not included in source–sink index calculations.
We furthermore calculated settlement symmetry between different regions for each species (Calabrese & Fagan, 2004). They calculated the forward settlement proportion, i.e., the symmetry of settlers from a specific settlement site (s) originating from an observed inception site (o), by scaling the number of successful settlers from site o settling at site s to each and every successful settlers originating from site o. Forward symmetry can live represented as Pso = Sos∕∑So. They furthermore calculated rearward settlement proportion, or the symmetry of settlers from a specific inception site (o) observed at settlement site (s), by scaling the number of settlers observed at site s originating from site o to each and every settlers observed at site s. The rearward symmetry can live represented as Pos = Sos∕∑Ss.Graph-theoretic analysis
To quantify connections between sites, they applied graph theory to population connectivity (Treml et al., 2008; Holstein, Paris & Mumby, 2014). Graph theoretic analysis is highly scalable and can live used to examine fine-scale networks between reef sites up to broad-scale analyses between islands or archipelagos, mapping to both local and regional management needs. It furthermore allows for both network- and site-specific metrics, enabling the comparison of connectivity between species and habitat sites as well as highlighting potential multi-generational dispersal corridors. Graph theory furthermore provides a powerful tool for spatial visualization, allowing for rapid, intuitive communication of connectivity results to researchers, managers, and the public alike. This type of analysis can live used to model pairwise relationships between spatial data points by breaking down individual-based output into a series of nodes (habitat sites) and edges (directed connections between habitat sites). They then used these nodes and edges to examine the relative import of each site and dispersal pathway to the greater pattern of connectivity around Moloka‘i, as well as differences in connectivity patterns between species (Treml et al., 2008; Holstein, Paris & Mumby, 2014). They used the R package ‘igraph’ to examine several measures of within-island connectivity (Csardi & Nepusz, 2006). Edge density, or the symmetry of realized edges out of each and every feasible edges, is a multi-site measure of connectivity. Areas with a higher edge density hold more direct connections between habitat sites, and thus are more strongly connected. They measured edge density along and between the north, south, east, and west coasts of Moloka‘i to examine feasible population structure and degree of exchange among the marine resources of local communities.
The distribution of shortest path length is furthermore informative for comparing overall connectivity. In graph theory, a shortest path is the minimum number of steps needed to connect two sites. For example, two sites that exchange larvae in either direction are connected by a shortest path of one, whereas if they both share larvae with an intermediate site but not with each other, they are connected by a shortest path of two. In a biological context, shortest path can correspond to number of generations needed for exchange: sites with a shortest path of two require two generations to beget a connection. average shortest path, therefore, is a descriptive statistic to estimate connectivity of a network. If two sites are unconnected, it is feasible to hold infinite-length shortest paths; here, these eternal values were eminent but not included in final analyses.
Networks can furthermore live broken in connected components (Csardi & Nepusz, 2006). A weakly connected component (WCC) is a subgraph in which each and every nodes are not reachable by other nodes. A network split into multiple WCCs indicates separate populations that accomplish not exchange any individuals, and a big number of WCCs indicates a low degree of island-wide connectivity. A strongly connected component (SCC) is a subgraph in which each and every nodes are directly connected and indicates a high degree of connectivity. A region with many small SCCs can attest high local connectivity but low island-wide connectivity. Furthermore, component analysis can identify slice nodes, or nodes that, if removed, burst a network into multiple WCCs. Pinpointing these slice nodes can identify potential famous sites for preserving a population’s connectivity, and could inform predictions about the impact of site loss (e.g., a large-scale coral bleaching event) on overall connectivity.
On a regional scale, it is famous to note which sites are exporting larvae to, or importing larvae from, other sites. To this end, they examined in-degree and out-degree for each region. In-degree refers to the number of inward-directed edges to a specific node, or how many other sites provide larvae into site ‘A’. Out-degree refers to the number of outward-directed edges from a specific node, or how many sites receive larvae from site ‘A’. Habitat sites with a high out-degree seed a big number of other sites, and attest potentially famous larval sources, while habitat sites with a low in-degree reckon on a limited number of larval sources and may therefore live contingent on connections with these few other sites to maintain population size. Finally, betweenness centrality (BC) refers to the number of shortest paths that pass through a given node, and may therefore attest connectivity pathways or ‘chokepoints’ that are famous to overall connectivity on a multigenerational timescale. BC was weighted with the symmetry of dispersal as described in the preceding section. They calculated in-degree, out-degree, and weighted betweenness centrality for each region in the network for each species.
As with the source–sink index, they did not embrace sites on islands other than Moloka‘i in their calculations of edge density, shortest paths, connected components, slice nodes, in- and out-degree, or betweenness centrality in order to focus on within-island patterns of connectivity.Results Effects of biological parameters on fine-scale connectivity patterns
The species-specific parameters that were available to parameterize the dispersal models substantially influenced final output (Fig. 2). The symmetry of successful settlers (either to Moloka‘i or to neighboring islands) varied widely by species, from 2% (Panulirus spp.) to 25% (Cellana spp.). Minimum pelagic duration and settlement success were negatively correlated (e.g., an estimated −0.79 Pearson correlation coefficient). Species modeled with batch spawning at a specific moon facet and/or time of day (Cellana spp., P. meandrina, and C. ignoblis) displayed slightly higher settlement success than similar species modeled with constant spawning over specific months. On a smaller scale, they furthermore examined average site-scale local retention, comparing only retention to the spawning site versus other sites on Moloka‘i (Fig. 2). Local retention was lowest for Caranx spp. (<1%) and highest for O. cyanea and P. sexfilis (8.1% and 10%, respectively).Figure 2: Summary statistics for each species network. Summary statistics are displayed in order of increasing minimum pelagic larval duration from left to right. Heatmap colors are based on normalized values from 0–1 for each analysis. Successful settlement refers to the symmetry of larvae settled out of the total number of larvae spawned. Local retention is measured as the symmetry of larvae spawned from a site that settle at the same site. Shortest path is measured as the minimum number of steps needed to connect two sites. Strongly connected sites refers to the symmetry of sites in a network that belong to a strongly connected component. live of value dispersal distance is measured in kilometers from spawn site to settlement site.
We measured network-wide connectivity via distribution of shortest paths, or the minimum number of steps between a given two nodes in a network, only including sites on Moloka‘i (Fig. 2). O. cyanea and P. sexfilis showed the smallest shortest paths overall, signification that on average, it would buy fewer generations for these species to demographically bridge any given pair of sites. Using maximum shortest path, it could buy these species three generations at most to connect sites. Cellana spp. and P. meandrina, by comparison, could buy as many as five generations. Other medium- and long-dispersing species showed relatively equivalent shortest-path distributions, with trevally species showing the highest live of value path length and therefore the lowest island-scale connectivity.
The number and size of weakly-connected and strongly-connected components in a network is furthermore an informative measure of connectivity (Fig. 2). No species in their study group was broken into multiple weakly-connected components; however, there were species-specific patterns of strongly connected sites. O. cyanea and P. sexfilis were the most strongly connected, with each and every sites in the network falling into a single SCC. Cellana spp. and P. meandrina each had approximately 60% of sites included in a SCC, but both demonstrate fragmentation with seven and six SCCs respectively, ranging in size from two to 22 sites. This SCC pattern suggests low global connectivity but high local connectivity for these species. Medium and long dispersers showed larger connected components; 70% of parrotfish sites fell within two SCCs; 40% of P. porphyreus sites fell within two SCCs; 70% of C. strigosus sites, 55% of C. melampygus sites, and 40% of Panulirus sites fell within a single SCC. In contrast, only 26% of C. ignoblis sites fell within a single SCC. It is furthermore famous to note that the lower connectivity scores observed in long-dispersing species likely reflect a larger scale of connectivity. Species with a shorter PLD are highly connected at reef and island levels but may demonstrate weaker connections between islands. Species with a longer PLD, such as trevally or spiny lobster, are likely more highly connected at inter-island scales which reflects the lower connectivity scores per island shown here.Figure 3: Dispersal distance density kernels. Dispersal distance is combined across species by minimum pelagic larval duration (PLD) length in days (short, medium, or long). Most short dispersers settle close to home, while few long dispersers are retained at or near their spawning sites.
Minimum PLD was positively correlated with live of value dispersal distance (e.g., an estimated 0.88 Pearson correlation coefficient with minimum pelagic duration loge-transformed to linearize the relationship), and dispersal kernels differed between species that are short dispersers (3–25 days), medium dispersers (30–50 days), or long dispersers (140–270 days) (Fig. 3). Short dispersers travelled a live of value distance of 24.06 ± 31.33 km, medium dispersers travelled a live of value distance of 52.71 ± 40.37 km, and long dispersers travelled the farthest, at a live of value of 89.41 ± 41.43 km. However, regardless of PLD, there were essentially two peaks of live of value dispersal: a short-distance peak of <30 km, and a long-distance peak of roughly 50–125 km (Fig. 3). The short-distance peak largely represents larvae that settle back to Moloka‘i, while the long-distance peak largely represents settlement to other islands; the low point between them corresponds to deep-water channels between islands, i.e., unsuitable habitat for settlement. Median dispersal distance for short dispersers was substantially less than the live of value at 8.85 km, indicating that most of these larvae settled relatively close to their spawning sites, with rare long-distance dispersal events bringing up the average. Median distance for medium (54.22 km) and long (91.57 km) dispersers was closer to the mean, indicating more even distance distributions and thus a higher probability of long-distance dispersal for these species. Maximum dispersal distance varied between ∼150–180 km depending on species, except for the spiny lobster Panulirus spp., with a PLD of 270 d and a maximum dispersal distance of approximately 300 km.Settlement to Moloka‘i and other islands in the archipelago
Different species showed different forward settlement symmetry to adjacent islands (Fig. 4), although every species in the study group successfully settled back to Moloka‘i. P. meandrina showed the highest percentage of island-scale local retention (82%), while C. ignoblis showed the lowest (7%). An average of 74% of larvae from short-dispersing species settled back to Moloka‘i, as compared to an average of 41% of medium dispersers and 9% of long dispersers. A big symmetry of larvae furthermore settled to O‘ahu, with longer PLDs resulting in greater proportions, ranging from 14% of O. cyanea to 88% of C. ignoblis. Moloka‘i and O‘ahu were the most commonly settled islands by percentage. Overall, settlement from Moloka‘i to Lana‘i, Maui, Kaho‘olawe, and Hawai‘i was more or less lower. Larvae of every species settled to Lana‘i, and settlement to this island made up less than 5% of settled larvae across each and every species. Likewise, settlement to Maui made up less than 7% of settlement across species, with P. meandrina as the only species that had no successful paths from Moloka‘i to Maui. Settlement to Kaho‘olawe and Hawai‘i was less common, with the exception of Panulirus spp., which had 16% of each and every settled larvae on Hawai‘i.Figure 4: Forward settlement from Moloka’i to other islands. Proportion of simulated larvae settled to each island from Moloka‘i by species, organized in order of increasing minimum pelagic larval duration from left to right.
We furthermore examined coast-specific patterns of rearward settlement symmetry to other islands, discarding connections with a very low symmetry of larvae (<0.1% of total larvae of that species settling to other islands). Averaged across species, 83% of larvae settling to O‘ahu from Moloka‘i were spawned on the north shore of Moloka‘i, with 12% spawned on the west shore (Fig. S4). Spawning sites on the east and south shores contributed <5% of each and every larvae settling to O‘ahu from Moloka‘i. The east and south shores of Moloka‘i had the highest average percentage of larvae settling to Lana‘i from Moloka‘i, at 78% and 20% respectively, and to Kaho‘olawe from Moloka‘i at 63% and 34%. Of the species that settled to Maui from Moloka‘i, on average most were spawned on the east (53%) or north (39%) shores, as were the species that settled to Hawai‘i Island from Moloka‘i (22% east, 76% north). These patterns attest that multiple coasts of Moloka‘i hold the potential to export larvae to neighboring islands.
Temporal settlement profiles furthermore varied by species (Fig. 5). Species modeled with moon-phase spawning and relatively short settlement windows (Cellana spp. and C. ignoblis) were characterized by discrete settlement pulses, whereas other species showed settlement over a broader age of time. Some species furthermore showed distinctive patterns of settlement to other islands; their model suggests specific windows when long-distance dispersal is possible, as well as times of year when local retention is maximized (Fig. 5).Figure 5: Species-specific temporal recruitment patterns. Proportion densities of settlement to specific islands from Moloka‘i based on day of year settled, by species. Rare dispersal events (e.g., Maui or Lana‘i for Cellana spp.) loom as narrow spikes, while broad distributions generally attest more common settlement pathways. Regional patterns of connectivity in Moloka‘i coastal waters
Within Moloka‘i, their model predicts that coast-specific population structure is likely; averaged across each and every species, 84% of individuals settled back to the same coast on which they were spawned rather than a different coast on Moloka‘i. Excluding connections with a very low symmetry of larvae (<0.1% of total larvae of that species that settled to Moloka‘i), they institute that the symmetry of coast-scale local retention was generally higher than dispersal to another coast, with the exception of the west coast (Fig. 6A). The north and south coasts had a high degree of local retention in every species except for the long-dispersing Panulirus spp., and the east coast furthermore had high local retention overall. Between coasts, a high symmetry of larvae that spawned on the west coast settled on the north coast, and a lesser amount of larvae were exchanged from the east to south and from the north to east. With a few species-specific exceptions, larval exchange between other coasts of Moloka‘i was negligible.Figure 6: Coast-by-coast patterns of connectivity on Moloka‘i. (A) average rearward settlement symmetry by species per pair of coastlines, calculated by the number of larvae settling at site s from site o divided by each and every settled larvae at site s. Directional coastline pairs (Spawn > Settlement) are ordered from left to right by increasing median settlement proportion. (B) Heatmap of edge density for coast-specific networks by species. Density is calculated by the number of each and every realized paths out of total feasible paths, disregarding directionality.
We furthermore calculated edge density, including each and every connections between coasts on Moloka‘i regardless of settlement symmetry (Fig. 6B). The eastern coast was particularly well-connected, with an edge density between 0.14 and 0.44, depending on the species. The southern shore showed high edge density for short and medium dispersers (0.16–0.39) but low for long dispersers (<0.005). The north shore furthermore showed relatively high edge density (0.20 on average), although these values were smaller for long dispersers. The west coast showed very low edge density, with the exceptions of O. cyanea (0.37) and P. sexfilis (0.13). Virtually each and every networks that included two coasts showed lower edge density. One exception was the east/south shore network, which had an edge density of 0.10–0.65 except for Cellana spp. Across species, edge density between the south and west coasts was 0.12 on average, and between the east and west coasts was 0.04 on average. Edge density between north and south coasts was particularly low for each and every species (<0.05), a divide that was especially several in Cellana spp. and P. meandrina, which showed zero realized connections between these coasts. Although northern and southern populations are potentially weakly connected by sites along the eastern ( P. meandrina) or western (Cellana spp.) shores, their model predicts very little, if any, demographic connectivity.
To explore patterns of connectivity on a finer scale, they pooled sites into regions (as defined in Fig. 1) in order to anatomize relationships between these regions. Arranging model output into node-edge networks clarified pathways and regions of note, and revealed several patterns which did not follow simple predictions based on PLD (Fig. 7). Cellana spp. and P. meandrina showed the most fragmentation, with several SCCs and low connectivity between coasts. Connectivity was highest in O. cyanea and P. sexfilis, which had a single SCC containing each and every regions. Medium and long dispersers generally showed fewer strongly connected regions on the south shore than the north shore, with the exception of C. strigosus. P. porphyreus showed more strongly connected regions east of Kalaupapa but lower connectivity on the western half of the island.Figure 7: Moloka’i connectivity networks by species. Graph-theoretic networks between regions around Moloka’i by species arranged in order of minimum pelagic larval duration. (A–D) Short dispersers (3–25 days), (E–G) medium dispersers (30–50 days), and (H–J) long dispersers (140–270 days). Node size reflects betweenness centrality of each region, scaled per species for visibility. Node color reflects out-degree of each region; yellow nodes hold a low out-degree, red nodes hold a medium out-degree, and black nodes hold a high out-degree. Red edges are connections in a strongly connected component, while gray edges are not fraction of a strongly connected component (although may still picture substantial connections). Edge thickness represents log-transformed symmetry of dispersal along that edge.
Region-level networks showed both species-specific and species-wide patterns of connectivity (Fig. 8). With a few exceptions, sites along the eastern coast—notably, Cape Halawa and Pauwalu Harbor—showed relatively high betweenness centrality, and may therefore act as multigenerational pathways between north-shore and south-shore populations. In Cellana spp., Leinapapio Point and Mokio Point had the highest BC, while in high-connectivity O. cyanea and P. sexfilis, regions on the west coast had high BC scores. P. meandrina and C. strigosus showed several regions along the south shore with high BC. For Cellana spp. and P. meandrina, regions in the northeast had the highest out-degree, and therefore seeded the greatest number of other sites with larvae (Fig. 8). Correspondingly, regions in the northwest (and southwest in the case of P. meandrina) showed the highest in-degree. For O. cyanea and P. sexfilis, regions on the western and southern coasts showed the highest out-degree. For most species, both out-degree and in-degree were generally highest on the northern and eastern coasts, suggesting higher connectivity in these areas.Figure 8: Region-level summary statistics across each and every species. Betweenness centrality is a measure of the number of paths that pass through a inescapable region; a high score suggests potentially famous multi-generation connectivity pathways. In-degree and out-degree advert to the amount of a node’s incoming and outgoing connections. Betweenness centrality, in-degree, and out-degree hold each and every been normalized to values between 0 to 1 per species. Local retention is measured as the symmetry of larvae that settled back to their spawn site out of each and every larvae spawned at that site. Source-sink index is a measure of net export or import; negative values (blue) attest a net larval sink, while positive values (red) attest a net larval source. White indicates that a site is neither a strong source nor sink. Gray values for Cellana spp. denote a want of suitable habitat sites in that particular region.
Several species-wide hotspots of local retention emerged, particularly East Kalaupapa Peninsula/Leinaopapio Point, the northeast point of Moloka‘i, and the middle of the south shore. Some species furthermore showed some degree of local retention west of Kalaupapa Peninsula. While local retention was observed in the long-dispersing Caranx spp. and Panulirus spp., this amount was essentially negligible. In terms of source–sink dynamics, Ki‘oko‘o, Pu‘ukaoku Point, and West Kalaupapa Peninsula, each and every on the north shore, were the only sites that consistently acted as a net source, exporting more larvae than they import (Fig. 8). Kaunakakai Harbor, Lono Harbor, and Mokio Point acted as net sinks across each and every species. Puko‘o, Pauwalu Harbor, and Cape Halawa were either weak net sources or neither sources nor sinks, which corresponds to the high levels of local retention observed at these sites. Pala‘au and Mo‘omomi acted as either weak sinks or sources for short dispersers and as sources for long dispersers.
Only four networks showed regional cut-nodes, or nodes that, if removed, burst a network into multiple weakly-connected components (Fig. S5). Cellana spp. showed two cut-nodes: Mokio Point in northwest Moloka‘i and La‘au Point in southwest Moloka‘i, which if removed isolated Small Bay and Lono Harbor, respectively. C. perspicillatus, and S. rubroviolaceus showed a similar pattern in regards to Mokio Point; removal of this node isolated Small Bay in this species as well. In C. ignoblis, loss of Pauwalu Harbor isolated Lono Harbor, and loss of Pala‘au isolated Ilio Point on the northern coast. Finally, in Panulirus spp., loss of Leinaopapio Point isolated Papuhaku Beach, since Leinapapio Point was the only larval source from Moloka‘i for Papuhaku Beach in this species.Figure 9: Connectivity matrix for larvae spawned on Kalaupapa Peninsula. Includes larvae settled on Molokaí (regions below horizontal black line) and those settled on other islands (regions above horizontal black line), spawned from either the east (E) or west (W) coast of Kalaupapa. Heatmap colors picture rearward proportion, calculated by the number of larvae settling at site s from site o divided by each and every settled larvae at site s. White squares attest no dispersal along this path. The role of Kalaupapa Peninsula in inter- and intra-island connectivity
Our model suggests that Kalaupapa National Historical Park may play a role in inter-island connectivity, especially in terms of long-distance dispersal. Out of each and every regions on Moloka‘i, East Kalaupapa Peninsula was the single largest exporter of larvae to Hawai‘i Island, accounting for 19% of each and every larvae transported from Moloka‘i to this island; West Kalaupapa Peninsula accounted for another 10%. The park furthermore contributed 22% of each and every larvae exported from Moloka‘i to O‘ahu, and successfully exported a smaller percentage of larvae to Maui, Lana‘i, and Kaho‘olawe (Fig. 9). Kalaupapa was not marked as a cut-node for any species, signification that plenary population breaks are not predicted in the case of habitat or population loss in this area. Nevertheless, in their model Kalaupapa exported larvae to multiple regions along the north shore in each and every species, as well as regions along the east, south, and/or west shores in most species networks (Figs. 9 and 10). The park may play a particularly famous role for long-dispersing species; settlement from Kalaupapa made up 18%–29% of each and every successful settlement in Caranx spp. and Panulirus spp., despite making up only 12% of spawning sites included in the model. In C. strigosus, S. rubroviolaceus, and C. strigosus, Kalaupapa showed a particularly high out-degree, or number of outgoing connections to other regions, and West Kalaupapa was furthermore one of the few regions on Moloka‘i that acted as a net larval source across each and every species (Fig. 8). Their study has furthermore demonstrated that different regions of a marine protected locality can potentially fulfill different roles, even in a small MPA such as Kalaupapa. Across species, the east coast of Kalaupapa showed a significantly higher betweenness centrality than the west (p = 0.028), while the west coast of Kalauapapa showed a significantly higher source–sink index than the east (p = 2.63e−9).Figure 10: Larval spillover from Kalaupapa National Historical Park. Site-level dispersal to sites around Moloka‘i from sites in the Kalaupapa National Historical Park protected area, by species. (A–D) Short dispersers (3–25 days), (E–G) medium dispersers (30–50 days), and (H–J) long dispersers (140–270 days). Edge color reflects symmetry of dispersal along that edge; red indicates higher symmetry while yellow indicates lower proportion. Kalaupapa National Historical Park is highlighted in light green. Discussion Effects of biological and physical parameters on connectivity
We incorporated the distribution of suitable habitat, variable reproduction, variable PLD, and ontogenetic changes in swimming competence and empirical vertical distributions of larvae into their model to increase biological realism, and assess how such traits impact predictions of larval dispersal. The Wong-Ala et al. (2018) IBM provides a highly supple model framework that can easily live modified to incorporate either additional species-specific data or entirely new biological traits. In this study, they included specific spawning seasons for each and every species, as well as spawning by moon facet for Cellana spp., P. meandrina, and C. ignoblis because such data was available for these species. It proved difficult to obtain the necessary biological information to parameterize the model, but as more data about life history and larval deportment become available, such information can live easily added for these species and others. Some potential additions to future iterations of the model might embrace density of reproductive-age adults within each habitat patch, temperature-dependent pelagic larval duration (Houde, 1989), ontogenetic-dependent behavioral changes such as orientation and diel vertical migration (Fiksen et al., 2007; Paris, Chérubin & Cowen, 2007), pre-competency period, and larval habitat preferences as such information becomes available.
In this study, they hold demonstrated that patterns of fine-scale connectivity around Moloka‘i are largely species-specific and can vary with life history traits, even in species with identical pelagic larval duration. For example, the parrotfish S. rubroviolaceus and C. perspicillatus demonstrate greater connectivity along the northern coast, while the goatfish P. porphyreus shows higher connectivity along the eastern half of the island. These species hold similar PLD windows, but vary in dispersal depth and spawning season. Spawning season and timing altered patterns of inter-island dispersal (Fig. 5) as well as overall settlement success, which was slightly higher in species that spawned by moon facet (Fig. 2). While maximum PLD did loom play a role in the probability of rare long-distance dispersal, minimum PLD appears to live the main driver of average dispersal distance (Fig. 2). Overall, species with a shorter minimum PLD had higher settlement success, shorter live of value dispersal distance, higher local retention, and higher local connectivity as measured by the amount and size of strongly connected components.
The interaction of biological and oceanographic factors furthermore influenced connectivity patterns. Because mesoscale current patterns can vary substantially over the course of the year, the timing of spawning for inescapable species may live faultfinding for estimating settlement (Wren et al., 2016; Wong-Ala et al., 2018). Intermittent ocean processes may influence the probability of local retention versus long-distance dispersal; a big symmetry of larvae settled to O‘ahu, which is more or less surprising given that in order to settle from Moloka‘i to O‘ahu, larvae must cross the Kaiwi Channel (approx. 40 km). However, the intermittent presence of mesoscale gyres may act as a stabilizing pathway across the channel, sweeping larvae up either the windward or leeward coast of O‘ahu depending on spawning site. Likewise, in their model long-distance dispersal to Hawai‘i Island was feasible at inescapable times of the year due to a gyre to the north of Maui; larvae were transported from Kalaupapa to this gyre, where they were carried to the northeast shore of Hawai‘i (Fig. S6). preliminary analysis furthermore suggests that distribution of larval depth influenced edge directionality and size of connected components (Fig. 7); surface currents are variable and primarily wind-driven, giving positively-buoyant larvae different patterns of dispersal than species that disperse deeper in the water column (Fig. S7).Model limitations and future perspectives
Our findings hold several caveats. Because fine-scale density estimates are not available for their species of interest around Moloka’i, they assumed that fecundity is equivalent at each and every sites. This simplification may lead us to under- or over-estimate the strength of connections between sites. want of adequate data furthermore necessitated estimation or extrapolation from congener information for larval traits such as larval dispersal depth and PLD. Since it is difficult if not impossible to identify larvae to the species flush without genetic analysis, they used genus-level larval distribution data (Boehlert & Mundy, 1996), or lacking that, an estimate of 50–100 m as a depth layer that is generally more enriched with larvae (Boehlert, Watson & Sun, 1992; Wren & Kobayashi, 2016). They furthermore estimated PLD in several cases using congener-level data (see Table 1). While specificity is example for making informed management decisions about a inescapable species, past sensitivity analysis has shown that variation in PLD length does not greatly impact patterns of dispersal in species with a PLD of >40 days (Wren & Kobayashi, 2016).
Although their MITgcm current model shows annual consistency, it only spans two and a half years chosen as neutral state ‘average’ ocean conditions. It does not span any El Niño or La Niña (ENSO) events, which understanding wide-scale sea-surface temperature anomalies and may therefore impress patterns of connectivity during these years. El Niño can hold a particularly strong impact on coral reproduction, since the warm currents associated with these events can lead to severe temperature stress (Glynn & D’Croz, 1990; Wood et al., 2016). While there has been runt study to date on the effects of ENSO on fine-scale connectivity, previous work has demonstrated increased variability during these events. For example, Wood et al. (2016) showed a reduce in eastward Pacific dispersal during El Niño years, but an increase in westward dispersal, and Treml et al. (2008) showed unique connections in the West Pacific as well as an increase in connectivity during El Niño. While these effects are difficult to predict, especially at such a small scale, additional model years would increase aplomb in long-term connectivity estimations. Additionally, with a temporal resolution of 24 h, they could not adequately address the role of tides on dispersal, and therefore did not embrace them in the MITgcm. Storlazzi et al. (2017) showed that tidal forces did impress larval dispersal in Maui Nui, underlining the import of including both fine-scale, short-duration models and coarser-scale, long-duration models in final management decisions.
We furthermore restrict their model’s scope geographically. Their goal was to determine whether they could resolve predictive patterns at this scale apropos to management. Interpretation of connectivity output can live biased by spatial resolution of the ocean model, since complicated coastal processes can live smoothed and therefore impact larval trajectories. To restrict this bias, they focused mainly on coastal and regional connectivity on scales greater than the current resolution. They furthermore used the finest-scale current products available for their study area, and their results demonstrate general agreement with similar studies of the region that employ a coarser resolution (Wren & Kobayashi, 2016) and a finer resolution (Storlazzi et al., 2017). Also, while learning of island-scale connectivity is famous for local management, it does disregard potential connections from other islands. In their calculations of edge density, betweenness centrality and source-sink index, they included only settlement to Moloka‘i, discarding exogenous sinks that would warp their analysis. Likewise, they cannot predict the symmetry of larvae settling to other islands that originated from Moloka‘i, or the symmetry of larvae on Moloka‘i that originated from other islands.
It is furthermore famous to note scale in relation to measures of connectivity; they await that long-dispersing species such as Caranx spp. and Panulirus spp. will demonstrate much higher measures of connectivity when measured across the whole archipelago as opposed to a single island. The cut-nodes observed in these species may not actually burst up populations on a big scale due to this inter-island connectivity. Nevertheless, cut-nodes in species with short- and medium-length PLD may indeed trace famous habitat locations, especially in terms of providing links between two otherwise disconnected coasts. It may live that for inescapable species or inescapable regions, stock replenishment relies on larval import from other islands, underscoring the import of MPA selection for population maintenance in the archipelago as a whole.Implications for management
Clearly, there is no single management approach that encompasses the breadth of life history and deportment differences that impact patterns of larval dispersal and connectivity (Toonen et al., 2011; Holstein, Paris & Mumby, 2014). The spatial, temporal, and species-specific variability suggested by their model stresses the necessity for multi-scale management, specifically tailored to local and regional connectivity patterns and the suite of target species. Even on such a small scale, different regions around the island of Moloka‘i can play very different roles in the greater pattern of connectivity (Fig. 8); sites along the west coast, for example, showed fewer ingoing and outgoing connections than sites on the north coast, and therefore may live more at risk of isolation. Seasonal variation should furthermore live taken into account, as mesoscale current patterns (and resulting connectivity patterns) vary over the course of a year. Their model suggests species-specific temporal patterns of settlement (Fig. 5); even in the year-round spawner O. cyanea, local retention to Moloka‘i as well as settlement to O‘ahu was maximized in spring and early summer, while settlement to other islands mostly occurred in late summer and fall.
Regions that demonstrate similar network dynamics may benefit from similar management strategies. Areas that act as larval sources either by symmetry of larvae (high source–sink index) or number of sites (high out-degree) should receive management consideration. On Moloka‘i, across each and every species in their study, these sources fell mostly on the northern and eastern coasts. Maintenance of these areas is especially famous for downstream areas that depend on upstream populations for a source of larvae, such as those with a low source–sink index, low in-degree, and/or low local retention. Across species, regions with the highest betweenness centrality scores fell mainly in the northeast (Cape Halawa and Pauwalu Harbor). These areas should receive consideration as potentially famous intergenerational pathways, particularly as a means of connecting north-coast and south-coast populations, which showed a want of connectivity both in total number of connections (edge density) and symmetry of larvae. Both of these connectivity measures were included because edge density includes each and every connections, even those with a very small symmetry of larvae, and may therefore embrace rare dispersal events that are of runt relevance to managers. Additionally, edge density comparisons between networks should live viewed with the caveat that these networks accomplish not necessarily hold the same number of nodes. Nevertheless, both edge density and symmetry demonstrate very similar patterns, and embrace both demographically-relevant common connections as well as rare connections that could influence genetic connectivity.
Management that seeks to establish a resilient network of spatially managed areas should furthermore account the preservation of both weakly-connected and strongly-connected components, as removal of key cut-nodes (Fig. S5) breaks up a network. Sites within a SCC hold more direct connections and therefore may live more resilient to local population loss. trust should live taken to preserve breeding populations at larval sources, connectivity pathways, and cut-nodes within a SCC, since without these key sites the network can fragment into multiple independent SCCs instead of a single stable network. This exercise may live especially famous for species for which they estimate multiple small SCCs, such as Cellana spp. or P. meandrina.
Kalaupapa Peninsula emerged as an famous site in Moloka‘i population connectivity, acting as a larval source for other regions around the island. The Park seeded areas along the north shore in each and every species, and furthermore exported larvae to sites along the east and west shores in each and every species except P. meandrina and Cellana spp. Additionally, it was a larval source for sites along the south shore in the fishes C. perspicillatus, S. rubroviolaceus, and C. strigosus as well as Panulirus spp. Western Kalaupapa Peninsula was one of only three regions included in the analysis (the others being Ki‘oko‘o and Pu‘ukaoku Point, furthermore on the north shore) that acted as a net larval source across each and every species. Eastern Kalaupapa Peninsula was particularly highly connected, and was fraction of a strongly connected component in every species. The Park furthermore emerged as a potential point of connection to adjacent islands, particularly to O‘ahu and Hawai‘i. Expanding the spatial scale of their model will further elucidate Kalaupapa’s role in the greater pattern of inter-island connectivity.
In addition to biophysical modeling, genetic analyses can live used to identify persistent population structure of relevance to managers (Cowen et al., 2000; Casey, Jardim & Martinsohn, 2016). Their finding that exchange among islands is generally low in species with a short- to medium-length PLD agrees with population genetic analyses of marine species in the Hawaiian Islands (Bird et al., 2007; Rivera et al., 2011; Toonen et al., 2011; Concepcion, Baums & Toonen, 2014). On a finer scale, they predict some flush of shoreline-specific population structure for most species included in the study (Fig. 6). Unfortunately, genetic analyses to date hold been performed over too broad a scale to effectively compare to these fine-scale connectivity predictions around Moloka‘i or even among locations on adjacent islands. These model results warrant such small scale genetic analyses because there are species, such as the coral P. meandrina, for which the model predicts transparent separation of north-shore and south-shore populations which should live simple to test using genetic data. To validate these model predictions with this technique, more fine-scale population genetic analyses are needed.Conclusions
The maintenance of demographically connected populations is famous for conservation. In this study, they contribute to the growing corpse of work in biophysical connectivity modeling, focusing on a region and suite of species that are of relevance to resource managers. Furthermore, they demonstrate the value of quantifying fine-scale relationships between habitat sites via graph-theoretic methods. Multispecies network analysis revealed persistent patterns that can befriend define region-wide practices, as well as species-specific connectivity that merits more individual consideration. They demonstrate that connectivity is influenced not only by PLD, but furthermore by other life-history traits such as spawning season, moon-phase spawning, and ontogenetic changes in larval depth. high local retention of larvae with a short- or medium-length PLD is consistent with population genetic studies of the area. They furthermore identify regions of management importance, including West Kalaupapa Peninsula, which acts as a consistent larval source across species; East Kalaupapa Peninsula, which is a strongly connected region in every species network, and Pauwalu Harbor/Cape Halawa, which may act as famous multigenerational pathways. Connectivity is only one piece of the puzzle of MPA effectiveness, which must furthermore account for reproductive population size, long-term persistence, and post-settlement survival (Burgess et al., 2014). That being said, their study provides a quantitative roadmap of potential demographic connectivity, and thus presents an efficacious tool for estimating current and future patterns of dispersal around Kalaupapa Peninsula and around Moloka‘i as a whole.Supplemental Information Current patterns in the model domain.
Current direction and velocity is displayed at a depth of 55 m below sea surface on (A) March 31st, 2011, (B) June 30th, 2011, (C) September 30th, 2011, and (D) December 31st, 2011. Arrowhead direction follows current direction, and u/v velocity is displayed through arrow length and color (purple, low velocity, red, high velocity). Domain extends from 198.2°E to 206°E and from 17°N to 22.2°N. The island of Moloka‘i is highlighted in red.Subset of validation drifter paths.
Drifter paths in black and corresponding model paths are colored by drifter ID. each and every drifter information was extracted from the GDP Drifter Data Assembly seat (Elipot et al., 2016). Drifters were included if they fell within the model domain spatially and temporally, and were tested by releasing 1,000 particles on the rectify day where they entered the model domain, at the uppermost depth layer of their oceanographic model (0–5 m).Selected larval depth distributions.
Modeled vertical larval distributions for Caranx spp. (left), S. rubroviolaceus and C. perspicillatus (middle), and P. porphyreus (right), using data from the 1996 NOAA ichthyoplankton vertical distributions data report (Boehlert & Mundy 1996).Coast-specific rearward settlement patterns by island
Proportion of simulated larvae settled to each island from sites on each coast of Moloka‘i, averaged across each and every species that successfully settled to that island.Regional cut-nodes for four species networks
Mokio Point and La‘au Point were cut-nodes for Cellana spp., Mokio Point was a cut-node for C. perspicillatus and S. rubroviolaceus, Pauwalu Harbor and Pala‘au were cut-nodes for C. ignoblis, and Leinaopapio Point was a cut-node for Panulirus spp.Selected dispersal pathways for Panulirus spp. larvae
500 randomly sampled dispersal pathways for lobster larvae (Panulirus spp.) that successfully settled to Hawai‘i Island after being spawned off the coast of Moloka‘i. Red tracks attest settlement earlier in the year (February–March), while black tracks attest settlement later in the year (April–May). Most larvae are transported to the northeast coast of Hawai‘i via a gyre to the north of Maui, while a smaller symmetry are transported through Maui Nui.Eddy differences by depth layer.
Differences in eddy pattern and strength in surface layers (A, 2.5 m) vs. profound layers (B, 55 m) on March 31, 2011. Arrowhead direction follows current direction, and u/v velocity is displayed through arrow length and color (purple, low velocity, red, high velocity). While big gyres remain consistent at different depths, smaller features vary along this gradient. For example, the currents around Kaho‘olawe, the small gyre off the eastern coast of O‘ahu, and currents to the north of Maui each and every vary in direction and/or velocity.
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