BIOLIntroduction to MarineF150 Biology Course Guide

An Independent Learning Course 3 credit hours

Developed and Graded by Susan DeLisa [email protected] University of Alaska Fairbanks

Copyright © 2012 University of Alaska Fairbanks (revised 2.28.13) The University of Alaska Fairbanks is an affirmative action/equal opportunity employer and educational institution.

iii Welcome & Introduction v v 3 7 97 19 37 53 67 79 87 75 105 115 129 137 141 145 151 161 123 157 ...... Table of of Table ...... Contents ...... The Human Factor Marine Mammals The Epipelagic Realm Life At The Top: The Netherworld: The Mesopelagic and Deep-Sea Realms On the Shelf: Continental Shelf Communities The Worlds: Between Two Intertidal Zone and Estuaries Microorganisms of and the Tree Cast of A Huge Life: Characters Tiny The Producers: Marine Autotrophs and Primary Production Marine Invertebrates—The Lower Phyla Marine Invertebrates—The Higher Phyla Marine and BirdsMarine Reptiles The Scope, History and Practice of The Benthic Environment The Environment Pelagic Organisms Cells, Biological Principles, & Ecosystems Course Outline and Requirements The Players: A Survey of Marine Organisms Marine of A Survey Players: The AllEcosystems It Together: Marine Putting Setting The Stage: Principles of Marine Science Marine of Principles Stage: The Setting Lesson 16 for Final Exam Request Appendix Lesson 12 Lesson 13 Lesson 14 Lesson 15 Lesson 11 for Exam #2 Request Three Part Lesson 7 Lesson 8 Lesson 9 Lesson 10 Part Two Part Lesson 5 Lesson 6 for Exam #1 Request Lesson 2 Lesson 3 Lesson 4 One Part Lesson 1 Preface Welcome Introduction iv BIOL F150: Introduction to marine biology v Welcome & Introduction - - - . The only . You will send your an your send will . You time limit of 4 hours , accordinginstructionsto the at found written assignment format,a with " to be filled out in duplicate, one copy to be sent one copy duplicate, in out " to be filled and a open-book Planning Schedule readingassignment to UAF eLearningto UAF and Distance Education , eacha with 16 lessons , one at the end ofend the at , one each of 3 partsthe of course,the instructions and are exams the taking insertedfor

3 exams

Thecourse includes swersassignments written to the work your "Survival be gradedwill websitetheir your or in returnedand one week. Therewithin Handbook." Usually, UAF to are also appropriateat the Coursethis places in are Guide. Exams their environments. In Part In environments. their we are introducedorganisms,marine to the Two, frombacteria, largestto the smallest the In whales. PartThree, organismsthese we visit habitatsmajor marine the where homes, their in organismsinteract each with other and their step we interactions backown our at look a critical for world.marine the with surroundings createto Finally, ecosystems. unique Course Organization and Requirements Course Organization Thiscourse is divided threeinto parts, eachinformation on the last, synthesizing building a into more and complete pic complex ture. Part One includes an overview of the scope and history of marine biologymethod, a and the scientific review of basic marine and basic biological and ecologicalnecessary principles ofand chemistry, the study physics for marine organismsgeology, and of this huge realm has been explored. Amazingly, we have better mapsof of realmhuge this our solar system, than in moon and other planets the been explored. has Amazingly, we have of At the same time, the floor! Marine biologynewfield, with exciting is an being made every discoveries day. theregrowing is concern about ofhealth the affectsthat how and ofoceans, health the the Theplanet. the future of , the we gain. knowledge the we use future,and indeed our own wisely system, and how complex this dependwe study well on how ing, since marine biologymarine since ing, reallynot is a separaterather but science, aspects of encompasses all biologyapplies as it to marine organisms nearlycover Marine environments environments. their and three quartersof areand planet the extremely varied, rangingfrom mangrove swampsand trenchesvents to volcanic pools, and tidal below the surfacemiles of lying the sea. The variety of diverse. and rawinhabiting life equally is food environments these depend for life and its ocean the on materials, We transportationbreathe we and recreation,the regulation and and even the oxygen of for fraction Earth's a tiny only climate. Yet, WELCOME WELCOME ofstudy scientific the 150, to Biol an Introductionsea. Thisthe in life Biology, to Marine mislead be can definition simple Welcome It also helps to have a systematic approach to studying. To get oriented, look at the list of oriented, look at the list get It also approachhelps to have a systematic To Objectives at the beginning to studying. of each end ofthe at assignment written the in questions and at the lesson, eachThen,lesson. readCoursethe material in the Guide read, reading the you As keeptextbook. the in assignment that by objectives the for followed lesson, and questions that for the backin lesson of show understanding. have a good Thenyou the material until To mind. Study answer the questions. your understandingyour ofwords, own ofthose not your and in answer must clearlymaterial, you the textbook, the and completely, and much new vocabulary to master. So, it is critical that you understand you that critical is it coursethe So, and much requirementsnew vocabulary a scheduleand plan to master. this, help with to complete the courseallow you and at the same time will you, be manageablethat will for on time. To 1 includes Lesson for a " assignment written the can record so you progress. you, your and monitor other to be keptand the completed assignment, by your with in materials that you may have to consult during exams will be this Courseare be this will exams the exams Although during to consult have textbook. may and your Guide you that materials open-book, rememberimportantis it that well-prepared,be to order in comfortably to time. allotted the in exam complete the Because of the vastand interdisciplinary scope, complexity nature of even at an introductory level, this marine biology, coursecovers of amount a huge information. are assignments new Reading there and extensive be many concepts, may vi BIOL F150: Introduction to marine biology Susan DeLisa received a B.S. in Biology from Antioch College, andM.S. Antioch aB.S. UAF. from andPhD. from inBiology Susan DeLisareceived inBiology degrees the instructorAbout "comealive." scribed inthiscourse and environments de canmaketheorganisms possible, especiallywhich thevideoclips, that you usethis site, if recommend sites. I strongly and links to related andvideoclips as flashcards, aids such summaries andquizzes, otherlearning with chapter a Hill) also offer (McGraw publishers The 978-0-07-352420-7). Guide and this Course are forthis course texts required The Required Textbooks enrollment. afteroneyear from given oradditional extensions will be NoIncomplete grades done and a passing grade). (generally, the work must have atleast half Handbook" in Semester-basedthe "Survival described students may for apply an incomplete IFthey the requirements meet completed. on work based agrade and will receive enrollment within one yearfrom All students must complete all coursework Extensions Incompletes and instructor. (orusual)assignmentfor isNOT thisclass. This arequired by credit writing a paper, passing, extra then it may by bepossible toearn the is borderline upon approval astudent's grade If Extra Credit following scale: tothe will assigned according be points). Final up to150 grades exams exam for your is worth (each 45% grade, forof 55% points. 1000 Written assignments account amaximum of out of points earned, Your of is calculated as the percentage grade Grading System andDistanceEducation. toUAF eLearning or at907-474-4011between8:00amand8:00pmAlaskaTime.Submitallwork you needhelp,you may contacttheinstructor,If other references. Guideor Susan DeLisa, [email protected] Course 78–79% 80–81% 82–87% 88–89% 90–91% 92–97% 98-100% ======C+ B– B B+ A– A A+ Marine Biology Marine Biology Marine 59–61% 62–67% 68–69% 70–71% 72–77% below 59% 9th Ed., by 9thEd., Peter Huber(ISBN andMichael Castro Website at www.mhhe.com/castrohuber9e ======F D– D D+ C– C - vii Welcome & Introduction : If are the semester time requirements student, follow you a semester-based book provided with your : Remember, we assume you will take up to a year to finish this course.this to finish take up to a year will we assume you At a baretake threeit should minimum, : Remember, you will experience the fascinationofexperience will you and appreciate marine life and marine environments their amazing diversityand and the great a better understanding, need for and better management, of these relationships complex interrelationshipscomplex better understanding of world your this course) beyond UAF Writing Center and Computer Lab offers Writing UAF telephone subject via free fax and student in any or to any writing tutoring Internet. the over information Students can call 907-474-5314 for on how to fax a paper and have it tutored the over telephone, or engage in an interactive Internet session. Both services are free. Off-Campus Library Services through Rasmuson Library the UAF can be arranged This calling 1-800-478-5348. by unit was set up to serve who students UAF to appropriate do not have access information resources in their town or village. 3) you will recognize will 3) you activities how human how the sea influencesinfluence our world, marine life and marine environments, 2) you will grasp will 2) you underlying the basic principles diversitythis a (and be for able to apply these principles and complexity 1) • • Let's begin to explore marine biology!

Goals of this Course of Goals The of goals this course are that: SEMESTER-BASED be accepted lessons cannot ifpurchase. turned Remember, in all at once. written assignment, be realisticassignment, written and try per up to two lessons arranged otherwise Unless to pace yourself. instructor, your with week should be submitted. YEAR-LONG months to complete a course. Instructor permission (in writing) must be obtained to finish in an accelerated time frame. stafffacultyand do everythingwill a lackof you, help to possible part your on planning emergencyan constitute does not for the staff and faculty! Please plan ahead! date planned of enter a date at least two weeks beforeexam,” Opposite “final your course completion. If are you a graduating beforeone month at least exam graduation.final to take the plan each submitting dates for planned entering When senior, Complete the Planning Schedule on the following pages and submit one copy (with your name and contact information) name and contact your (with with one copy pages and submit Schedule Planning Complete the following the on firstyour the appropriate lesson. Put course deadline for bottom of completion at the the schedule. If are you a graduating check counselor to find out the date gradesall your with are due. Ifsenior, are you graduating other personal or any have imperativeis it instructor deadlines, your to write first you that your on informand lesson ofhim/her the Although needs. your Planning Schedule Other Important Information Important Other viii BIOL F150: Introduction to marine biology ix Welcome & Introduction Grade ______Received Feedback Feedback ______Actual Date Sent ______Submit Date to ______

14 15 16 12 13 4 5 6 7 8 9 10 11 1 2 3 Grade must be received ______. by I began this courseon ______.

Final Exam Lesson Lesson Lesson Exam 2 Lesson Lesson Lesson Lesson Lesson Exam 1 Lesson Lesson Lesson Lesson Lesson Lesson Lesson Lesson

DEADLINE: START: Planning Schedule (Keep copy.) this BIOL F150: Introduction to marine biology x xi Welcome & Introduction Submit Date to ______. ______. ______. ______Grade must be received ______. by I began this courseon ______.

Lesson 12 Lesson 13 Lesson 14 Lesson 15 Lesson 16 Final Exam Lesson 7 Lesson 8 Lesson 9 Lesson 10 Lesson 11 Exam 2 Lesson 2 Lesson 3 Lesson 4 Lesson 5 Lesson 6 Exam 1 Lesson 1 DEADLINE: CONTACT PHONE: CONTACT START: Planning Schedule first with your copy (Submit this lesson.) NAME: STUDENT E-MAIL ADDRSS: xii BIOL F150: Introduction to marine biology 1 Welcome & Introduction

Setting The Stage: Stage: The Setting Principles Marine of Science Part One Part

3 Lesson 1: Scope, History and Practice of Marine Biology . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson) for completing the work for required in this course.

schedule Reading Assignment Reading Written Assignment 9th Ed.). eLearning & Distance Education. Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Plan a realistic Gain an historical perspectiveexploration on human and use ofand its the sea resources. Learn about technologies the marine world. studying for Understand how the scientific method works, science is an open-ended and why process. Appreciate the difficulties inherent the marine world, in investigating and the extent to which it remains unexplored. Start thinking about some of the fundamental concepts and questions in Marine Biology. the information of pages Read this lesson in the following for this Complete the Biology Complete the More information, including interactive flashcards, quizzes, videoclips can be and links, 1. 2. 3. 425–432) 1–16) and Chapter 19 (pp. C&H, Chapter 1 (pp. 3. 4. 5. 6. 1. 2. Reading assignment Reading steps for completing the lesson the completing steps for Marine Biology may be simply defined as the scientific study of as the scientific study defined Marine Biologybe simply may living things in saltwater in this brief And here for environments. simplicity ends, definition we have just introduced three vast, and open-ended subjects. complex this Lesson of Objectives and Practice of and of Practice Marine Biology Scope, History History Scope, 1 BIOL F150: Introduction to marine biology 4 M terrestrial world? New organisms are being discovered all being discovered are organisms New world? terrestrial to ourown compared themarineworld, of ignorance greater to our Or is this disparity due in part biodiversity? Earth’s contain only really one-eighth of on Earth, space living ity of major the vast includes which Doesthe sea, worldwide. fied species havebeenidenti million 1.9 marine andterrestrial and life-styles. As many as forms playing incredibly diverse (not counting dis been described, bacteria) have organisms marine species of andunknown. About 240,000 equally vast may be realm unexplored andlargely immense, varied things ( living of variety The discovered? to be waiting are oceanworlds What new tem! in our solarsys surfaces mapped poorly the more one of is the Earth) of the surface (most of the seafloor In fact, to be explored. the ocean remains than of 95% ments, more marineenviron of variety astaggering have discovered Yet,canoes andhot springs, amongothers. scientists while vol anddeep-sea reefs, coral swamps, mangrove marshes, polaroceans, aswell as salt seas andice-covered tropical turquoise darkness, perpetual of realms and vast waters rine environments. Saltwater ma of diversity arich alsohelp to shape rivers, of outflow andtides,and the currents wind,waves, of effects varying conditions. andchemical The light, pressure temperature, umn and along the sea floor, of span a huge range which col zones,both in the water numerous this continuum are Along km 11(7miles) below sealevel. asnearly as deep by lapped the tides to trenches shorelines extending from space, its living of 95% and fill about the Earth, of 70% than subjects.ended environments covermore Saltwater complex andopen- vast, three we havejust introduced The Scope, History Practice of Biology and Marine And here simplicity ends, for in this brief definition simplicity ends,for in And here this brief environments. things in saltwater living tific study of may besimplyMarine Biology definedasthe scien (Spying ontheSea:ShipsandSonar, ScubaandSubmarines,Satellites andSensors) biodiversity include sunlit surface sunlit surface include ) residing in this ) residing • ------all the life that Earth contains, which brings us to the third brings us to the third contains, which all the life that Earth still, we may know never surprising to these questions. More Amazingly,discovered? answers not haveconclusive we do speciesyet tobe thetotalnumberof magnitude, of order toan Can weestimate,even known to us, oronly afraction? already biodiversity Earth’s Is most of level. microscopic welook,especially atthe everywhere catalogued) or even faster,the time(much than they described, in canbe fact, cess. Scientific knowledge and is is sub constantly growing we don’t know! So, science is an ongoing,pro open-ended we doknow may know - andwhat palein comparison towhat weknow, that we don’t the more words, werealize the more In other questions than it answers. often brings up more science Good information. synthesizing andviewing ways of and equipment,by techniques new new of development continuallyby beingopened vistas are the andnew search, is continuallyby being added information on-going New re scientific research. at many levels,of oversight, the careful thescientific method andby by theuseof exerted control knowledge is anything but stable, though, for all the quality- aspossible.Scientific knowledgeis asreliable of store to our contributed information pieceof thateach toensure process called the dure, conducted by science) followingare basic proce the same of studying(as well marinelifeas thosein all otherareas to approaches the different a globalscale.However,all of on photosynthetic marine organisms of the productivity andcomputerizedmathematical models togauge imagery dolphins, orusing satellite captive of agroup tion skills of byin marinebacteria the laboratory,testing the communica performed conversions volve investigating specific chemical lifein the sea could in about disciplines. Learning different study, fieldof many broad including is a very Marine Biology “scientific study.” MarineBiology: element in our definition of scientific method . We. usethis time-tested ------5 Lesson 1: Scope, History and Practice of Marine Biology ------, exterminatingforms unique of life, species of ? Our own future these be strongly will influenced by As you learn about Marine Biology, and come to a more learnyou As about Marine Biology, complete understandingof themes, essential these some ofconsider also raise:they that questions the What kinds of marine environments remain to be ex plored? How much of Earth's biodiversity is still un and what known types to us, yet of are marine to be life by shaped been organisms marine have How discovered? physi and communities biological their with interactions shaped be to continue they will how and environments, cal sci future and current will How interactions? future by marine the of understanding our change research entific on activities our of impacts the are what Finally, world? ma change impacts those will how and ocean, world the conditions affect and life, marine and environments rine globally depend and will changes, part in we understand on how well world,the marine knowledge. we use that and how wisely 7) As you will learn will 7) As you relied always have lesson, humans this in next the In resources, ocean and its on the ways. many in learn will you two lessons, ocean that waters and the sea floor regulatealso our atmosphere and climatedrive and weather-relatedsuch phenomena as drought, hur flooding, ricanes and tornadoes. Theyare geo responsible many for logical processes including and events, mountain-building, earthquakes,and even tsunamis of the position volcanoes, globe! on the The continents the resources, ocean and its turn,in are increasingly subjected to impacts from human aremarine habitats destroying and marine life We activities. and at an alarming scale on a huge rate, eventhe causing extinction some in cases even before have been discovered. they This perhapsis most important the "take-home message" from this course. ------geo . As a resultof these , the family tree family , the of relation ecosystem and Tree ofTree Life (included as an AppendixCourse to this . Thiscourse an evolutionary emphasizes community evolution Guide) and to the Guide) ships among all living things. 6) The most importantconceptunifying biology in that is of perspective,order in staggeringthe put to help diversity of marine organismsrespect both with context, into the to logic timescale interactions, energy flowscycle and nutrients from organism to organism, and between organisms and the environment. level,basic most its At ofmade up is life ofexchanges these energy and matter. 5) Another recurrent5) Another coursethis theme in interdepenthe is dence oforganismsall eachwith physi their with and other Eachliving integralan environments. cal is thing part of a biological ever they look, scientists using newusing look, scientists ever they techniques are finding microscopic organismsunimaginedin numbers and diver performingfact,ocean. the In in functions vital many sity, microorganismsappear diversemost abundant, to be the and important least known) forms (but of life on the planet. 3) Theinhabiting life extremely is also environments these diverse to us. unknown be mostly and may Much4) of see. Wher can eye the than smaller is life this revising knowledge. our 2) Theworldmarine ofcomprises most Earth'senviron is extremely varied,ments, and largely unexplored. number ofnumber "themes" repeated throughoutcourse,this im portant exploreideas to keep Marine Biology: as you mind in understandingour to key the Science, 1) of world,the is refiningand adding to, an open-ended process, continually ject to continual refinement, continual to ject even and revision, occasional perpetually building, under construc complex a huge like remodeled, being continually and tion occasional the with and reconstructiondemolition of some parts. It is a human endeavor and, as such,imperfect. This is firstthe is of a BIOL F150: Introduction to marine biology 6 on time! quired the Complete materialsyouonline ateLearning.uaf.edu received. andinthecourse a Include thesubject.) hasnoknowledgescribing/explaining tosomeonewho of "explain"or "describe" something,don't just so - list. do (To de thatyouimagine are and showyour clear understanding, be own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 9. 8. 7. 6. 5. 4. 3. 2. 1. ), and keep a copy for yourself—check this schedule periodically to make sure you are on track to finish the course tofinish the course on track you periodically are sure tomake this schedule copy a ), andkeep foryourself—check Explain why the study of life in the oceans is important. (give three reasons). three (give lifeintheoceansis important. Explain why the study of theSeatreaty. IstheUnitedStatesamongthem? theLawof Unionhavesignedandratified and theEuropean 2012,163countries disputes. Asof of usage, andtheresolution sovereignty, rightsof of matters and addressing themarineenvironment andits resources, andequitable usage of for oceans, theconservation providing world's the the useof for regulating theSeaestablished aframework 1982UnitedNationsConvention ontheLawof The tellhowyou briefly (Inotherwords, experiment.) wouldconductacontrolled results? donot"confuse"the thatothervariables seaweed,howwouldyou makesure of rate thegrowth intensity increases light it? Whenconductinganexperiment increasing thatmightaffect totestwhether othervariables examples of functioning. three thatmay Whatare anorganism's influence isonevariable itssurroundings of temperature The tellhowitmightbetested.Define"scientifictheory."briefly hypothesis atestable and your from own experience, anexampleDefine "testablehypothesis." of give Preferably any subject). (concerning reasoning anddeductive inductive of own experience,oneexample each give ontime.From your ontime,thenImightdeducethat you willfinishthecourse that studentscompletethiscourse impression Ihavethegeneral If weekuntilyou finish thecourse. intwoassignmentsevery thatyouclude willturn tocon intwoassignmentsthisweekandnext reasoning week,Imightuseinductive you turn If themdirectly. andobserving inperson being there or by atadistance,without thatallowscientists to study scubadiving. Describefour marineorganisms technologies inthelaboratory, attheshore, onshipsand mannedsubmarines, marineorganisms observe Scientists candirectly sonar,What dothefollowing standfor: acronyms scuba,AUV, ROV, COOL,UNCLOS, EEZ? marinescience. fundamentally advanced thisvoyage ways inwhich Describethree isbasedonitsresearch. oceanography theocean,andmodern study of theHMS voyage of The usedtheoceananditsresources. Describe four ways thathumanshavetraditionally cover sheet cover , not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , Planning Schedule Written Assignment for 1 Written Lesson Assignment and submit the written assignment to UAF eLearning & Distance Education according to directions found to directions & Distance Education according andsubmit the written assignment to UAF eLearning (Please make your completeandconcise) answers Challenger on p. ix of this Course Guide. Return one copy of your with this assignment ( schedule onecopy of Guide.Return this Course onp.ix of (1872-1876) was the first worldwide expedition dedicatedtothescientific worldwide thefirst (1872-1876)was • brief and clear , but, also complete . For example, if asked to asked For. example, if in yourin - re - - 7 Lesson 2: Benthic Environment - - Marine Course Guide.

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Written Assignment Reading Assignment Reading 9th Ed.). Complete the eLearning & Distance Education. Online Learning at the publisher's be found Center: www.mhhe.com/castrohuber9e Understand the concept of a worldbe able ocean and to identify its principal geo graphical and gulfs. regions: bays seas, the major ocean basins, ofHave a basic knowledge the internal structure of the Earth and the characteris tics of the continental and oceanic crusts. Gain an historical perspective of the exploration of the sea floor and appreciate the to whichdifficulties and the extent inherent these environments, they studying in remain unexplored. ofBe familiar with the "anatomy" the sea floor—its profiles, zones and major features. Understand the process of plate tectonics and how it transforms the Earth's surface and shapes the ocean environment. the information of pages Read this lesson in the following for this Complete the Biology More information, including interactive flashcards, quizzes, videoclips can and web-links, 1. 2. 3. 18-38) C&H: Chapter 2 (pp. 2. 3. 4. 5. 1. steps for completing the lesson the completing steps for assignment Reading Objectives of this Lesson of Objectives environment The is vast, benthic environment and largely unexplored. complex Submarine topography is dramatic,not only dynamic. it is also extremely The Benthic Benthic The 2 BIOL F150: Introduction to marine biology 8 T correspond to the peach seed, pit and flesh, respectively.pit andflesh, seed, to thepeach correspond Earth's The peach. toa might be compared ourmulti-layered planet,which of layer is the outermost crust dynamic tremely processes. The the Earth's of distinct parts have collected water, andgeologically chemically but are that not simply hollows in the Earth ocean basins are The the benthic environment. about asyou read for reference, a your is textbook, there of At the ocean. the back of regions specific geographically or identified as arms and gulfsare Pacific Basin and South Indian Basin), and many seas,bays specified (e.g.,major basins, smaller basins the Northwest are The intomajor 4 is classified theEarth, coveringof 71% The otherandwith their environments. with each their interactions principles governing and general organisms, living istics of basic character by reviewing 4 completes our preparation, environment. Lesson conditions in the pelagic and chemical benthic environment, and in Lesson 3 wewill study physical the of andprocesses the physicalwe explore features Here, zones. specifichabitat more into several is divided ronments column floor, andthe The pelagic. tats two general oceanincludes dynamic environment. The this and continue to shape that created and theprocesses the ocean sons,of wewill study the physical characteristics fold-out map of the world ocean theworld fold-out mapof Pacific , the environments where organisms live: benthic and live: organisms , the environments where world ocean world some background knowledge.In thenext twoles some background we need this course, 2 of sea their home, in Part thatmakethe To formeeting the organisms prepare above the bottom. Each of these two general envi thesetwogeneral of Each the bottom. above , benthic Atlantic pelagic , the interconnected expanse of seawater seawater expanse of , the interconnected inner core ("depth") habitat is in that on or the sea habitat ("depth") , ("sea") habitat comprises the ("sea") habitat Indian crust and The Benthic Environment , outer core Arctic Oceans Arctic , and are subject to ex , andare . Keep this map open this map Keep . and . Within the basins mantle water water habi • - - - - - .

less dense, so the continents float higher on the underlying higherontheunderlying so thecontinents less dense, float granite thicker basalt called rock adark-colored tively thin. It is mainlyof composed things. skin. It holds the oceans, continents fuzzy peach and all living to the corresponding thinnest layerThe by is the crust, far surveys and chart the nation's coasts. The first hydrographic hydrographic first the nation's coasts. The and chart surveys to conduct in 1807 Congress and the U.S., Jefferson established by Thomas was President The sounding by this immense, unseen surface, sounding! of the ocean and decipherthe shape of ing to plumb the depths try it was, how difficult during the process. Imagine current the boat andline being subjected to movement by wind and to makeonesounding, It could take hours arms. outstretched to thedistance between aman's equivalent rope ments of in measured was hit Depth bottom. aship until the weight from lowering a weighted hemp rope the Old English "sund" meaning sea/water/swimming), from bymade were measurements Early in the ocean. depth of surement using theseafloor about learned We have known simply to observe. because it is so difficult complex and still un largely benthic environment is vast, The Mapping History of Seafloor Exploration and in this lesson. about wewill learn theplanet,which of surface in plate tectonics, involvingthe dynamic processes the entire significantly also figure crust ences between the two typesof Density differ is submerged. the oceanic crust mantle while Survey of theCoast of Survey , which is rich in iron and therefore quite dense. The quite The dense. andtherefore in iron is rich , which continental crust continental Oceaniccrust , which is rich in silicon and aluminum and therefore in silicon and aluminum and therefore is rich , which sounding (not related to the word "sound" but to theword (not related , underlying the ocean basins, is rela , underlying consists largely of lighter-colored lighter-colored consists of largely , the oldest scientific agency in the oldestscientific , agency hydrographic bathymetry fathoms (bathymetric) , 6-ft incre , 6-ft , the mea , ------9 Lesson 2: Benthic Environment ------). and (Global Challeng GPS during World World during , which produce echo sounding sonar voyage, hydrographic voyage, Seismic reflection side scan sonar Challenger's (Light Detection And Ranging)And Detection (Light systems and LIDAR expedition, which expedition, happened.have otherwise not might vide seamless coverageseamless vide ofelevation depthand between measurementsbathymetric also allows land and sea. LIDAR and rugged along complex to be made safely shorelines. sonar and preciseSide scan measurements of the relative strengthof reflectedalso can be used to determine signals the textureof and composition sandy, the sea floor (rocky, as depth.as well etc.) muddy, Around time of the surveyingwas greatlyreplacement the advanced by of hemp ropeof wire with invention the lines, wireline the sounding machine,and deep-sea was It anchoring. truly revolution development the of by ized, however, the previous saw in 1.7) Wars(C&H Fig. lesson II. We I and beam ofa single sonar using how activeused to sonar is measure depththe of the sea floor ( hydrographic mapping is carried using out primarily Today, sonar multibeam a broad swathof 1). The echo (Fig. soundings navigationglobal satellite the System), system, Positioning accurately determinesthe geographic ofposition the sur oflocation and the vessel vey eachEarth.on the sounding The of amount huge data generated processedis and im addi In specialized computer systems. with aged digitally to sonar, tion ofpulses use emitted from laser light, aircraft and reflected offshoreon objects waterin and pro50 m, to to down we shall find them teeming with life." Thelife." animal with them teeming find we shall debate dredgingwas officiallyoperations ended by fromconducted 1867, Coast Sur In death). the 1867-1869 Forbes' (after prolific found deepas life vey 517 fathoms as off coast the of1868, and in the NorwegianMichaelscientist Florida, Sars documented 427 species broughtup from 450 fathoms off results,these InspiredScottish the ofcoast by the Norway. naturalistCharles Thomson Wyville of (a student Forbes') and Admiralty Society fund to Royal British the petitioned dredging operationsoff ofcoasts the and Ireland, Scotland productive1868 in and 1869. Many were hauls made well below 300 fathoms, including one at 4,289 m (2,346 fath deepest the oms), ever dredgeemphatically time, that at haul was It successes these hypothesis. azoic disprovingForbes' led to Thomson'sthat organization of the HMS er

------ex Terror and Challenger Erebus 29' W, the middle of 29' W, o 26' 17 and longitude S o to bring up the mud and stones ofto bring up the mud and stones the bed of the ocean, and other life fromand other dredgings as deep as 400 fathomsduring which1840-1843 their expedition, prompted to write, Ross contrary"although generalto the belief ofnaturalists, have I fromthat doubt no however greatbe able a depthmay we Forbes declared below 300 fathoms. Forbes exist life didn't that Sometimes referred we know it to to as the "Azoic Theory," announce waitingto be disproved. Forbes' be a hypothesis, startedment a debate 20 years,lasted for that despite the and crewfactRoss that ofup baskets full had hauled corals miles ofmiles rope and 12 ofmiles piano wire sailed as they nearly115,000 throughkm Arctic.the except oceans the all The presence of life on the deep-sea floor also was unknown naturalist1843, In British the mid-1800s. the until Edward make more deep-water includingsome deeper soundings, first.the than wasIt 1872-1876 the HMS firstthat pedition conducted and documented deep-sea a largeon soundings begin to scale enough giveto some characterizationto the floor of the world144 ocean, using portofa weight January On 76 lbs. reached 3, 1840, they 27 latitude BrazilOcean between Rio de Janeiro, Atlantic the South and sea sounded the they Namibia. Here, dead in calm weather, floorcrew at 2,425 fathoms(14,550 ft). Ross's on to went ing the 1839-1843 voyage ofthe 1839-1843ing voyage HMS toward sailed as they Antarctica, under the command of James Clarksearch in Ross, of A Magnetic South the Pole. wasline constructed on boardwas that 3600 fathomslong, intervalsat fitted swivelswith strongand sup to enough The deep-sea floor wasn't sounded until the mid-1800s. Thedeep-sea floor wasn'tmid-1800s. the until sounded believed)then, a bottomless ocean (as some Until was a been provenyet that hadn't false! Thetestable hypothesis firstdeep-water successful were soundings taken dur continental shelfcontinental breakslope (from and continental sound westernthe in ings North 1849), firstin the Atlantic known offMonterey (in Bay seafloor canyon the coast of California 1857)in and the first of indications and trenches. seamounts maps showed only the largestmapsfeatures only showed benthic shallow in patternswaters,coastal for looking painstakingly by detected They served soundings. among many haz identify to primarily earlyards its During to near-shore shipping. years, however, SurveyCoast the also was responsiblediscovery the for of the 10 BIOL F150: Introduction to marine biology National Oceanic and Atmospheric Administration, formed in formed National Oceanic andAtmospheric Administration, of is now a part agency), conservation the first in 1871, Fisheries (formed Commercial of and the Bureau to atmospheric science) dedicated agency the first in 1870, CoastSurvey,The with together the Weather (formed Bureau AUVs and ROVs have continued exploring the ocean floor. like vey vessels,submersibles sur of mantle. In addition to drilling vessels and afleet and betweenthe crust transfers massandenergy vestigate the sea floor,and in change, on climate seekinformation under Its mission is toexamine life deep effort. Japan-led by countries. 25 of led consortium byin 1983 OCP, involving a U.S.- the Ocean Drilling Program andplatetectonics. succeeded It was spreading seafloor of lenger out with the vessel carried Drilling Project ft beneath the 7,000 ocean floor.nearly as asdeep from the earth samplesof has pulled upcore (Fig. within the sea floor 1). meters of downtohundreds features pulses, subsurface detect ergy refraction seismic IOCP from 1968-1983, provided key evidence in support in keysupport evidence provided 1968-1983, from , the Integrated Ocean Drilling Program, aU.S.- OceanDrilling Program, and , the Integrated , echo sounding using explosive en echo , Alvin OCP was followed in 2003 was Figure 1. Hydrographic mappingtechniques. (C&HFig.and 1.10), DSDP Deep-sea drilling lmr Chal Glomar , the Deep Sea , the Deep NOAA , the - - - - and investigations have pieced together the broad outlines and investigations the broad together havepieced byone headed (including Ross) many searches years, 150 Erebus months later, Bay siting in Baffin afew After abrief Franklin. SirJohn Passage,of under thecommand plete theNorthwest in an attempttocom the Arctic toward north they turned 1845, hulls.with steam engines and iron-plated In May of vessels asbomb in the Royal outfitted Navy)were careers their long began these sailing ships (which to the Antarctic, AftertheRoss expedition exploration. toseafloor respect ror Epilogue foryou at is available toexplore that,however, of (Half mapped. has been comprehensively the sea floor 1900s. Still, of only 10% about and early 1800s the andscientists of byhave beenimagined theexplorers the sea floor, could not probably which of and processes the features have madeimmense strides in understanding on their information a wealth of and you theglobe, canfind around peditions and research NOAA Commerce. conducts extensive andexciting ocean ex of agencies under the 12 U.S. Department as oneof 1970 is legendary and, in a strange way, and,in a strange is legendary with comesfull circle and : The rest of the story of HMS of thestory of rest : The Terror were never seen again. Over the next seen again. never were oceanexplorer earth.google.com/ocean Erebus website. We website. and er Te !) - - - 11 Lesson 2: Benthic Environment , . ------The " in the " in and hot spots hot Titanic balanced herself (or tablemounts) are (or tablemounts) Mid-Atlantic Ridge Galapagos SpreadingGalapagos submarine canyons Trieste Guyots designer, AugustePiccard, de designer, ) burn) through there(although is Trieste's also paid a visit to the Challenger Deepto the Challenger 1995. also paid a in visit , partof east the in system mid-ocean ridge the mantle plumes mantle , among other films!) also descended into the Chal the into descended also films!) other , among Kaiko ocean's crustocean's where of columns rock hot from deepthe in earth( 36-37). About 50 hot spots have debate about C&H this: pp. Hot spots under partsbeen identified. of mid-ocean ridge the aresystem to have formed thought Azoresthe like islands 2.7) the on (C&H Fig. and Iceland Galapagosthe and the at Islands Center ment arement including known, offThe Canyon Hudson Submarine the coast of New the Grand than wider and longer Can deeper, is York of most With yon. sea floor the unexplored, still what formationsother amazing remain to be discovered? Processes Dynamic The dramatic displays sea flooronly not features,also un it dergoes continually processes. Sediment extremely dynamic rains and avalanches plains, abyssal down on the of rock and Lavasand sweep erupts through canyons. submarine from forming which volcanoes, cracks grow into the sea in floor, rise above Seamounts surface the seamounts. to become is erode and islands surface,below the lands, and sink away again seamounts. becoming flat-topped formerly seamounts, whose islands, tops became and wavesand wind subsided. Chains eroded as they by other groupingsof also form islands " over and 48 minutes, the ascent, 3 hoursand 48 minutes, Pic 15 and minutes. card, son ofthe scribed the touchdown "the thus: few the on delicately ofpounds the on rope guide lay that claim,token bottom, making ofname the in and hu science depthsultimate to the Chal oceans - the our all in manity, Theirfeat wasrepeated not Deep." lenger more for 50 than years,James Cameron until (director of Abyss 26, a submersible in Mar. designed, on he Deep, lenger 2012. descent took aboutHis 2 and a half hours, the ascent ("heckuva 70 minutes only ride!"), and live of updates the event were from posted on Twitter a yacht the accompanying The expedition. remote-controlled Japanese-built, submers ible other dramaticMany features ofenviron the benthic ------as Mauna Terror , also can , also Challenger Challenger , the longest vallongest , the seamounts , near Guam in the west , in near Guam , chaina colossal of volca . Her descent took 4 hours . It lies at depthslies . It of 3,000- Trieste central valley rift Mariana Trench abyssal abyssal plain . The (fittingly, Greek "deep for darkness")and (fittingly, , plunging to about, plunging 11 below sea km (7 mi) level. If , lying mostly submergedofmostly island on the , lying Hawai'i, is mid-ocean ridge system mid-ocean ridge trenches scaphe ("deep boat") than a mile of water. In 1960, a real-life "Voyage to the Bot the to ofa mile 1960, than In a real-life"Voyage water. tom of the Sea" place took when Jacques Piccard, a Swiss an Walsh, Don Navy Lieutenant U.S. and undersea explorer, Deepthe bathy in the Challenger descended into engineer, at erndeepestthe holds Pacific, the known, place Deep would Mt. it Everest were Deep, the Challenger tipped into ofout sink morebe coveredbase would and it's sight by Kea the world's 9,632 It rises mountain. tallest m (31,601 ft) from more base to summit, 2,500 than Everest. than ft taller The Earthplaces on lowest also are sea floor, on the found great depression, the on Earth.ley Individualgrow that volcanoes to 1,000 m called or more abovesea floor, the breakto enough high rise seamounts be seen. When the One suchvolcano, surface, become islands. they the encirclesthat mountains nic the globe, runningthrough all nearly 2.5). Occupying a third (C&H Fig. oceanbasins the oflargestthe is this geologic formation on ocean floor, the Earth.along the top of Running a is ridge the mid-ocean surface, the is 5,000 m (10,000-16,500 flattestthe and is ft) place planet, coveredon the hundreds,is that mud with even ofthousands, The vast of feet deep. expanse the abys features. The many brokenis sal plain most dramatic by is Dramatic Features Dramatic The sea floor can be described superlatives.in The most geographicextensive feature on Earth, 60% of covering its rationParks with Canada, side scan and mul has been using chartto ROV sonar and an tibeam sea floorthe search in of an effort in and the ships, to strengthenCanada's claim of sovereignty largeover portionsof a rapidly warmingArctic. of ofdemise the searchthe but Franklin the expedition, for 1992, In Canada HMS designated continues. the vessels Erebus undiscoveredonly (their Site Historic sucha National site). 2008, Hydrographic Since Canadian the Service, collabo in 12 BIOL F150: Introduction to marine biology running parallel with its axis. The blocks at the crest of the of at the crest blocks with its axis. The parallel running other,each along themid-oceanridge system, prominent are from have beendisplacedvertically thefracture ther sideof has fractured. crust with is littered floor a day. allthis seismic activity,earthquakes of the sea result As a four noes. Vailulu'u, for of example, experiences anaverage the mid-ocean ridges and other volca neartrenches, areas the ("shaking") activity. characterize Frequent earthquakes also issubject tomuch ridgefloor system, thesea vents,seamounts and themid-ocean duces hydrothermal ( In additiontoallthevolcanic activity million 6-8 years. completelyonceevery about itself cling in this way, crust recy theearth's passes through water (C&H Figs.It is and 2.27). thought 2.26 that all the ocean's called " build structures into chimney-like solidify and which dense clouds producing with minerals, these vents is laden emittedfrom Often the water surface. floor, up to the becomessuperheated,and is vented back in the sea downinto countless cracks seeps seawater when association with the mid-oceanridge systemandtrenches, occurin They theocean. chemistry, of physics and biology the of ourunderstanding these vents changed of covery dis Alvin. The in the with late 1970's the aid of discovered hotsprings called lifearound of holds astonishing oases systems.floor Even the deep-sea and mostcomplex marineeco the richest reefs, by coral fringed volcanic islands are Many tropical marine organisms. community of a thriving supports already Nafanua summit of alsois home todynamic activity. seafloor biological The The in Polynesian event culture. a highly revered would be born, island that happens, anew If growth. of rate its present within decades, the surface at cone could breach this new war, the Samoan goddessafter of Nafanua Named in 2001. this seamount since the last bathymetric measurements of within the crater m tall, cone, 300 had appeared that a new andROV, and deploying a submersible techniques they found lands,the sea floor. is rising from Using bathymetric mapping hotspot volcano, theVailulu'u seamount near the Samoan Is another anisland where of anticipatinghave been thebirth Ecuador. the coastof scientists Since 2005, Pacific off ern faults Normal faults Normal , or rifts , where the brittle oceanic , where , where the blocks onei the blocks where , hydrothermal vents hydrothermal volcanism black smokers ) thatpro seismic ------" - ,

mantle. into the downandrecycled is being pushed back sea floor time, called is process This growing). asyour fingernails the same rate onaconveyor belt(moving at about as if outward, spreads that seafloor to new the mid-ocean ridges, birth giving of is continually the mantle at the crest magma rising up from is constantly sea floor in motion. Molten the entire In fact, the ocean basins (Fig. through zigzags 2and C&H Fig. 2.5). it in themanyridge, placessothat offsetting it the axis of other horizontally. to occur perpendicular faults Transform faults mid-ocean ridge system also displaysThe many (Fig. riftvalley 2andC&HFig. line,the central 2.25). crest along the running a depression creating ridge havedropped, quakes. As a result of these plate interactions, oceanic crust oceanic crust these plate interactions, of quakes. As a result explosively, is eventually released which earth causing more and building tension catching faulting), transform other (i.e., past each involves two plates scraping a shearboundary (C&H Fig.byfied Fault2.15), the San Andreas in California calleda plateboundary type of way, in third a plates interact zones attrenches, third along a zonesatmid-oceanridges andthe subduction spreading Plate). In addition to the is being pushed into the Eurasian other (e.g. the Indian Plate each the Himalaya,against where as the plates buckle created are mountain ranges great collide, neither is subductedand continental bearing crust and melt onits way downinto the mantle.Whentwoplates Fig. up as the subducted 2.8), plate edgetobreak begins commonnearthese subduction zones (C&H volcanoes are and (C&HFig.Earthquakes 2.13). atrench also creating andpushes the other down, one overrides oceanic crust, (C&H Fig. plates both carry When the 2.12). two converging a pushing it into the mantle and creating anic crust, other, the dense oce overrides the lighter continental crust on the on oneplateedge andcontinental crust oceanic crust they collide with neighboring plates. When a collision involves zones, thespreading away from As theseplatesgrow 2.11). over the globe (C&HFig. like giant puzzle pieces arranged lithospheric duction subduction , where the two sides of the fracture slip past each slip past each the fracture the two sides of , where seafloor spreading Spreading zones Spreading

zones

plates at trenches, mark the boundaries of great great theboundariesof mark attrenches, is occurring at the trenches, where old where atthetrenches, is occurring , which partition the Earth's surface surface the Earth's partition which , atmid-oceanridges, and (C&HFig.At the same 2.10). shear boundary transform transform . Exempli trench sub - - - -

13 Lesson 2: Benthic Environment - - - - - Illustration by Christen Bouffard, UAF eLearning - - Te ("all Gond ("all sea"). ("all Pangaea Panthalassa , to the north,, to the and . rifts formedAs and flooded Laurasia Sinus Borealis and , to the south, filled with water from the Tethys Sea, waterfilled with , to the south, fromTethys the continents, separatingcontinents, Theand seas. oceans smaller rift dividing Pangaea into wana creating the NorthOcean. GondwanaAs Atlantic broke apart, offsplit India northward, and moved Indian the establishing from away America moved Ocean, and South Africa, creating BorealisOcean; the Sinus Atlantic the South shifted north unifying theoryunifying draws of it together evidence from geology, disparate branchesof seismology and paleon (e.g., science tology—the of study where to explain fossils) earth and why how ocean occur, volcanoes and mountain-building quakes, open and close,basins drift, and collide continents and how apart,split transforming continually the face of our planet. and crustthe all almost was probably sea floor, Originally, cyclesof spreadingwere and subduction rapid. By about years3 billion graniticrockago, sur the at up built had face to form numerousmicrocontinents, which eventually clumped together to formcore the of our modernconti Aroundyears220 million whennents. dinosaurs ago, ap peared,Earth the one supercontinent, held earth"),and one gigantic ocean, off bays Two enclosed Panthalassa enormous the seas, thys Sea Pangaea began to break apartseawater, with smaller into ------Figure 2. Hydrologic cycle. Hydrologic 2. Figure , whichoccurs subduction, during (tectonics means "to build") is the all-en is means "to build") (tectonics accretion lation oflation overwhelmingevidence. is achieved Often this with the aid of new technologies,of or the synthesis information from differentof All disciplines. factors these were involved developmentthe in of whichrevhas theory, plate tectonics our understandingolutionized of planet. The dynamic this counted in your textbook, is a great is textbook, of example your in counted as an science process—gradually ongoing refining,and oc and expanding leaps sweeping or huge making casionally alterations, our in understandingof the world. The face strong latter typically first,at opposition gaining after acceptanceaccumu the only Plate tectonics Plate compassing term describing the creation, defor movement, and destructionmation of through plates, lithospheric the cyclicthe creation, spreading ofsubduction and sea floor. the The story of how the theory ofdeveloped, plate tectonics re as sediments on the subducted plate areplate subducted the on sediments as scraped offand plastered plate. Lava onto the overiding eruptingfrom volca added continents. to the is material that way another is noes recycled, whilecrust continental up and mostly built being is endures. Therockscontinental oldest known are nearly 4 bil yearslion old, whereassea flooroldest the wasformed "just" a are Continents yearssome 200 million up by built ago. process called is relativelyand short-lived, thin being it is continually since 14 BIOL F150: Introduction to marine biology house effect toanenhanced part like these, duein large changes being felt, especially Global already in coastal areas. are level in sea these eustatic changes impactsof The rate. erating melting at anaccel are expanding andglaciers are waters havebeenincreasing.Warming temperatures and ocean decades, however, In recent thousand years. atmospheric little in the last few ago, hadrisen very andsealevel years had meltedby that iceage 10,000 from about theglaciers of ago.years Most 18,000 the about last ice age, the peak of todayft) higherthan at m (425 130 and sealevel is about it cools. when period, We now in an interglacial contracts are and increases, its the ocean also temperature expands when ods as iceandsnow, falls. During sealevel riods eustatic changes.tuations During also produce Climatic fluc aeustatic change. is an example of spreading ocean basins due to seafloor es. Change in the volume of called in global adjustments level, to sea toas referred motions, andfault are as well astoearthquakes processes, onalocal scale duetosuch surface theEarth's uplift of oroil.Subsidence or groundwater of orextraction rocks, of weathering and erosion glaciers, the melting of through orsediments accumulate. Unloading occurs is extruded, lava ice accumulates in glaciers, when Loading happens removed. weight or being added of masses sink or rise as aresult time.Land geologic ocean basins and continents,through between the the relationship also change Other factors today, platetectonics continues.granted of asthe process we takefor the arrangement from different be very again oceansandcontinents will of the configuration the future, Far East Africa. into of in the riftvalleys basin is forming ocean an "embryonic" basin, while broad ing a mature Red Seais a "juvenile" basin, on its wayThe to becom subducted. are sin, continuing to shrink as its borders Panthalassa)ba is a "declining" of Ocean (the remnant basins, still expanding Pacific by spreading. The seafloor Today, "mature" Oceans are Indian and Arctic the Atlantic, ago.million 200 years about these changes, from lustrates Fig.Ocean. the Arctic to become in your 2.17 textbook il , sea level rises again as glaciers melt. The volume of volumeof melt.The as glaciers rises again sea level , , when more of the Earth's water accumulates on land water theEarth's of more when , , will be discussed more in the next lesson. will discussed, more be isostatic changes. Other processes result result changes. Otherprocesses interglacial peri interglacial eustatic glacial pe chang green ------continental margin is covered by a thick wedge of sediment by wedgeof iscovered athick continental margin (C&HFig.The there 2.23). operating tectonic processes plate by is shaped the type of area in aparticular margin the continental floor. the deep-sea gin, toform of profile The the continental mar away from abyssal plain stretches The this slope, a the base of sloping steeply and a more the continental crust, the end of marks a is submerged, nental crust sloping sea floor the two main regions: (C&H Fig. all have They features 2.20). the same general basins share all theocean theglobe, similar ways around in floor the sea toshape work Because theseprocesses The Sea Floor Profile increasing darkness, cold and pressure. The The cold andpressure. darkness, increasing the mid-ocean ridge system.It is subject to of rise, andmuch the continental slope and m, including 4000 to about m deep) (usually 120-200 break the shelf zoneextends from "deep") The and currents. waves atlow tide, but submerged is exposed tosunlight, remains It below the intertidal. the continental shelf including break, sublittoral the tideis out. The and totheairwhen waves, exposed tothe sun and between the low and high tide marks, The organisms. of communities different supports zone, andeach in each differ tidal, bathyal, abyssal, and hadal zones (Fig. 3).Conditions sub the benthic environment: the intertidal, of divisions five are the ocean basins, there of to this profile Corresponding Benthic Habitat Zones rise. Ocean),agentle slope andapronounced the Arctic mi, in the caseof 620 km, or than 1,000 (more long shelf a edge,cur at aplate edge, then it has oris on a receding called an forms, anddynamic profile andasharp down into the trench, carried sediments are colliding and subduction is occurring, two platesare plate edge,where an advancing near curs acontinent oc thecontinent. When theedgeof off eroded passive margin passive continental slope continental shelf . The continental margin generally exhibits generally a gently continental margin . The active margin active ) zone runs from the low tide mark to the shelf totheshelf thelow tide mark from ) zone runs intertidal with the wedge of sediment forming a sediment forming with the wedge of continental margin bathyal . When a continent does not oc continental rise . When sediment accumulates at , where the edge of theconti the edge of where , shelf break shelf (or littoral (from the Greek bathys, the Greek (from ) zone is the area ) zone is the area that more or less or that more , and the will also form. will also form. abyssal subtidal deep- (from (from (or - - - - - Shelf Break Low Tide Tide Low Mark High Tide High Tide Mark Abyssal Hadal Baythal Subtidal Intertidal Benthic Habitat Zones Benthic Habitat

15 Lesson 2: Benthic Environment Shelf Break Low Tide Tide Low Mark High Tide High Tide Mark The bathyal, abyssal and hadal zones together make up the The abyssal bathyal, basin this in fill we will lesson, next the In "deep-sea floor." ofpelagicthe with zones benthic when habitatzones, we characteristics physical the study of pelagic the environment. - - Figure 3. Benthic habitat zones.. Benthic habitat 3. Figure Abyssal Hadal Baythal Subtidal (from the classical Hades, Greek under Intertidal

hadal sunlight and subject to extremeand subject sunlight pressure and cold, except at hydrothermal which vents, are habitatshottest the Earth.on eLearning UAF Bouffard, Christen by Illustration the Greek abyss, "bottomless") zone comprises the sea floor the zone comprises "bottomless") Greekthe abyss, fromand deep plains 4000 6000 to m, ocean the abyssal Thebasins. world)refers zone 6000 trenchbelow bottom m, the the to hadal zones are and without abyssal the Both completely es. 16 BIOL F150: Introduction to marine biology your textbook. of attheback theglossary guideandinyour textbook, andcheck thematerialinthiscourse re-read theseterms, any of themeaningof about unclear you're If for understanding. for thislesson.Doaself-check somekey terms Listed beloware • • • • • • • • • • • • • • • • • • seamount andguyot riftvalley central and EastPacific Rise mid-ocean ridge system,Mid-AtlanticRidge abyssal plain oozeandsiliceous calcareous ment lithogenous sedimentandbiogenous sedi sonar/echo soundingandLIDAR bathymetry/hydrography continental drift lithospheric plates andplate tectonicstheory andlithosphere asthenosphere basalt andgranite andcontinentalcrust oceanic crust inner core, outercore, mantle, andcrust Oceans Pacific, andSouthern Atlantic,Indian,Arctic, ocean world column water benthic andpelagic glossary terms - • • • • • • • • • • • • • • • • • seafloor spreading and spreading zone andspreading seafloor spreading fault andtransform fault normal fault/rift, volcanism andseismicactivity andblack smoker vent hydrothermal Deep MarianaTrench,trench, andtheChallenger hot spot/mantleplume submarine canyon bathyal, abyssal andhadal subtidal(sublittoral), (littoral), intertidal margin andactive margin passive slope andcontinentalrise continental break, continental shelf, shelf isostatic andeustatic andLaurasia Gondwana Pangaea andPanthalassa accretion anomalies magnetic shear boundary subduction andzone 17 Lesson 2: Benthic Environment -

in your in your . For example, if example, . For asked to complete , but also , but clear and brief

• : normal faults are cracks in the earth's crust where two sides are the and the Challenger Deep and the Challenger Terror transform faults (Please make your answers and concise) complete your make (Please and and submit the written assignment to UAF eLearning to UAF assignment written the and submit according Education & Distance to directions found and HMS Written Assignment LessonWritten 2 for active margins and passive margins asthenosphere and lithosphere Gondwana and Laurasia sediments and biogenous sediments lithogenous sonar and LIDAR Trieste Glomar Challenger and Mauna Kea Mariana Trench and Galapagos Islands guyots benthic and pelagic , ofnot those or Course textbook the Guide. Be

cover sheet h. i. j. d. e. f. g. a. b. c. in the development of the theory of Describe the pattern plate tectonics. observed they how it supports and explain ofthe hypothesis seafloor Describe two other kinds of spreading. observations of the sea floor around the mid- also support they why that scientists have made, and explain ocean ridge of the hypothesis seafloor spreading. page – on next assignment continued – Written Identify the two principal types of rock crusts. that distinguish the continental and oceanic Describe three other important differences continental and oceanic crust. between Observation is the currency of science. When scientists towed a magnetometer and plot above the mid-ocean ridge observed they ted the data on a map, a surprising pattern of a discovery magnetic role "stripes," a key that played Name the four major basins ofName the four the world ocean. Which continents separate which from oceans each other? displaced vertically, while in transform faults the sides are displaced horizontally. Along the mid-ocean ridge, normal the mid-ocean ridge, Along while in transformdisplaced vertically, faults the sides are displaced horizontally. faults are parallel to the axis while transform faults are perpendicularof to the axis the ridge. Define the following pairsDefine the following of also include but information. all the essential terms. Comparing/contrasting Be brief, the example: two terms For to do this. will often help you normal faults 3. 4. 2. 1. online at eLearning.uaf.edu and in the course received.online at eLearning.uaf.edu materials you own words understanding, be clearyour and show are imagineto you that explaining (To list. just do so - don't "describe" something, someone who of has no knowledge the subject.) Includea Study the information presented for this lesson until you feel that you have a good understanding. Write your answers your information the understanding. have a good Study Write feel that you presented you until lesson this for 18 BIOL F150: Introduction to marine biology 6. 5. ably expect tofindonyour trek.) Youcontinental margins. may you alsouseyour mightreason tocomeupwithsomegeologic features imagination of 2,andtheillustrations inC&HChapt. thetextbook, othermaps of (Usethefold-out attheback map this route. floor. anddeep-sea order,margins inchronological Describethesefeatures asyou wouldencounterthemalong onthecontinental you features mightseealongthispath,including inthislesson,which about you havelearned features 11specificseafloor (withlabels) Describeordiagram Portugal. andendingatthewestcoastof chushetts, ,Massa atCape theoceanfloor,Put onyour across scubagear andtakeawalk starting route alongadirect these boundaries. of ateach andshearboundaries. occurring themid-oceanridge Describetheprocesses system,trenches, of valley rift thecentral thelithosphericplatesare theboundariesof thatform crust theEarth's of majorfeatures three The - - 19 Lesson 3: Pelagic Environment - Marine Course Guide.

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). Climate Change" and "Ocean Acidification: The Other CO2 Problem") and p. 239 ("Will 239 ("Will and "Ocean Acidification: TheClimate Change" Other CO2 Problem") and p. There Be a Last Straw?") Online Learning at the publisher's be found Center: www.mhhe.com/castrohuber9e Appreciate the extraordinary properties chemical and physical of and be water, familiar cycle. with the hydrologic Be familiar with the chemical characteris composition of and the physical seawater, tics of the pelagic including environment, sea ice. Learnis structured how the ocean according to temperature, vertically, salinity and density. Understand the concept of thermohaline circulation and its impact on global climate patterns. the major surfaceRecognize currents of are how they the ocean, generated and regulatehow they climate, understanding and have a basic of waves and tides. Be able to classifypelagic the into habitat environment zones. Understand the significance of in the heat balance of the ocean the planet, and as a moderator of the greenhouse effect and global warming. the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education. More information, including interactive flashcards, quizzes, videoclips can and web-links, http://highered.mcgraw-hill.com/sites/0073028193/student_view0 1. 2. 3. 40–62), and "Special Report" The 231–237 ("Rolling Dice: C&H: Chapter 3 (pp. pp. 4. 5. 6. 7. 1. 2. 3. 98% ofwater the cycles that Earthon continually is also ocean. Water the in held is being circulated patterns complex ocean, the in within stronglythat influenceboth climate,and global marine environments affecting thus of all Earth's inhabitants. Water is a simple molecule with complex and extraordinarycomplex with molecule simple a is Water properties. the It is primary requirementofconstituent principal the and life for Close to livingthings. Reading assignment Reading steps for completing the lesson the completing steps for Objectives of this Lesson of Objectives environment The Pelagic Pelagic The 3 20 BIOL F150: Introduction to marine biology C Venus, tothe sun, has an atmosphere our neighbor closer and climate. atmosphere Earth's influence strongly in turn, in the ocean, occurring for life.Processes requirement mary it in a liquid state, the pri andkeeping this water retaining for responsible part in large are maintain our atmosphere, and that created the sun, and the complex processes from Our distance tions continue to bringsteamthesurface. have also contributed water,the Earth and volcanic erup Comets and meteorites striking to cool and harden. it began theplanet,as of surface ontheearly themolten rock from originated as steamemitted water This space. from viewed when blue is a predominantly sphere mi), our world (2.3 precipitating in the form of rain or snow. rain On land, this pre of in the form precipitating blown by are thewind overeventuallythesea, landor which clouds plants,then forms theleavesof from is transpired andland, the oceans, lakes, from rivers as itevaporates in the maining 5% participates re The is bound into rocks. surface on Earth's thewater of lifeonland.About 95% the distribution of determines largely water of availability things. living The principal constituent of for life and the requirement It is the primary properties. Watersimple moleculewith is a complex andextraordinary exists in the ground. occuratthe poles andpermafrost caps although ice at the surface, topersist cold for liquid water in the bydistant past, but water is now too eroded flowing tohavebeen appears landscape Martian The –60°C. of temperature andanaverage than ours thinner atmosphere boiled away eonsago.with a much On theothersideis Mars, it once heldwould have Any(800°F). 450°C liquid water house effect with a runaway times denser than Earth's, 100 71% by seawater at an average depth of 3.7 km 3.7 of depth by71% atanaverage seawater by covered water, its surface planet. With of 74% the water is really toits "Earth" name, Contrary producing a surface temperature of about about of temperature surface a producing The Pelagic environment hydrologic cycle hydrologic (Fig. 4), green • - - - - - patterns that strongly influence both marineenvironments influence that strongly patterns is continuallywithin in the ocean, complex beingcirculated is held in the ocean. Water water this cycling also of 98% continues.its way tothe ocean aswell, as the cycle Closeto finds and mostgroundwater tothesea, flow which into rivers drain waters melt, surface plants. Glaciers up by of the roots groundwater, to become taken and is again into the earth percolates builds glaciers, lakes andrivers, cipitation refills discovery that came out of the HMS that came out of discovery This salinity, ocean. of the world throughout regardless same, (C&H Table seawater orless the more remain 3.1) ions of ice (through ice of formation (through water of due totheremoval %). Salinity (or pptor in some placesmayhigher, be usually psu is 35 seawater salinity of average The cally equivalent. units) or or as asapercentage, be expressed 3.1. listed in C&H Table are in seawater ions present types of main The NaCl. (Cl-),in of the case dium (Na+) and chloride into their component electrically-charged in water ( nantly sodium chloride are the solutes in seawater Mostof erties. prop its many remarkable substance,oneof other natural kinds of more dissolved materials,3.5% orsolutes. Water candissolve and water 96.5% of on average, is composed, Seawater CompositionChemical inhabitants. Earth's all of thus affecting and globalclimate, ). However, the proportions of the major constituent themajor of ). However,theproportions Salinity rule of constant proportions of rule ppt precipitation (parts per thousand), all of which are numeri are which thousand), all of per (parts , the amount of salt dissolved in the water, may, the amount of solutes ), or lower due to the addition of water water ), or lower due to the addition of (i.e., is (i.e., a better NaCl , river runoff river - table salt), which salt), -table which Challenger psu is another important is another important evaporation or the or solvent (practical salinity (practical salts melting of melting of dissociate expedition. ) than any , predomi ions or the or : so : - - - -

21 Lesson 3: Pelagic Environment

. - - - - . Over photosyn bicarbonate ) is present) is at reacts chemi gas to be ab 2 2 2 gas exchange (CO calcium calcium carbonate , marine organisms, marine up take ) readily enter the sea surface 2 (O Carbon dioxide Carbon respiration oxygen ions, which ions, moreallows CO )and 2 (N they produce oxygen and absorb carbon dioxide. and absorb carbon dioxide. produce they oxygen carbonate decreasing salinity, decreasing temperature, and increasingdecreasing salinity, pressure. The primary influenceon gas concentrationsbelow the surface of the activityis of ocean, the however, marine organisms. During while and releaseduring carbon dioxide, oxygen thesis marine organismsMany also incorporate into carbon dioxide made of shells and skeletons til equilibrium is reached,is equilibrium til a processcalled wavesdue to and currentsTurbulence accelerates this process. gasesAs the most abundant the atmosphere, in nitrogen through gas exchange. concentrationits but 500 is very low concentrationsthe air, in greatertimes surface in waters,the most prevalentit making gas ocean. Thethe in more50 times ocean holds carbon atmosphere.the in than Thisbecause CO is incorporated becoming into water, with cally and sorbed sea surfaceat the throughgas Gas con exchange. centrationsvary different in regions and at different depths of the solubility the ocean. Generally, in gases increases with - - - - - , the Figure 4. Hydrologic cycle. Hydrologic 4. Figure Nutrients , chemicalsreleased releasefrom materials erosion Pollutants and Weathering

, as well as solids, dissolve in waterin dissolve are and solids, as , as well important Illustration by Christen Bouffard, UAF eLearning UAF Bouffard, Christen by Illustration interface,from moving to a lower concentration a higher un way into the ocean and can profoundly affectprofoundly ocean and can the into way life. marine Gases ofconstituents Gases diffuse across air-sea the seawater. from the sea floor, throughhydrothermal and subma vents fromsea floor, the from and volcanoes, rine whosevolcanoes land emissions rain ocean. the on down which humans by negatively affectofquality the envi the ronmentfertilizers(like fumes), also find their and exhaust the functioning and distribution of and distribution functioning the and life. Ions marine a varietyenter the ocean in other nutrients of (C&H ways 3.6). Fig. rockswhichand soil, are washedrivers into or picked by up and carriedwinds, also aresea. to the Chemicals released materials needed for life, includeneededand other for materials major ions the requiredwhich substances nearlyin occur may undetect The concentra critical, concentrationsable, yet seawater. in oftion can vary some nutrients greatly and the ocean in strongly influences concentration,nutrient as salinity, as well In addition to the major ions, small amounts of amounts small major In addition to the ions, virtually all other chemicalsseawater. in can be found 22 BIOL F150: Introduction to marine biology of that produced by human activities in the past 200 years. by that produced in years. the past 200 human activities of dioxide, carbon half about and storinghugequantities of by absorbing andglobalwarming, effect the greenhouse have beenmoderating ocean andits inhabitants The fect. warming cement production. andfrom burned), carbon ton of foreach dioxide produced carbon are tons 3½ of (about fossil fuels that we have been burning mous quantities of the enor carbondioxide from released of levels increasing this warmer. tobecome sphere Since the industrial revolution, causing the atmo the Earth, of the surface from radiated is a carbon dioxide the air. from Atmospheric carbon dioxide huge for absorbing quantities responsible of are ganisms shells carbonate and skeletons that they build, marine or photosynthesis, Also andthe through calcium we breathe. the oxygen that of half sible into the air about for releasing respon are their photosynthesis, marineorganisms Through sediments accumulated on theseafloor, contributing to theseshells have quantities of time,immeasurable eons of quired to raise the temperature of 1 gram of asubstance of 1gram of the temperature toraise quired is its unusually high solvent power. the most important its unsurpassed Oneof in addition to Water hasmany unusual physical properties, Temperature calcium carbonate). shells and skeletons of als to form like shellfish and cor marine organisms of ing the ability (e.g., processes limit and biological rocks) weathering of of both physical (e.g., the rate affect processes increasing continues. This ent trend the pres if further much industrial times andmay increase since pre- 30% about theoceanhasincreased of surface it more of centration in seawater, chemicals a higher con producing balance of dioxide carbon shifts the chemistry. water addition of The sea it has also altered the atmosphere, of ated warming carbondioxide has moder uptake of While the increased greenhouse effect greenhouse greenhouse gas greenhouse acidic is the result of this increasing greenhouse ef greenhouse this increasing of istheresult and forming carbonate rocks like limestone. carbonate rocks and forming hydrogen ions hydrogen . The hydrogen ion concentration near the hydrogen ion concentration . The heat capacity that prevents the escape of heat of the escape that prevents has been increasing as a result of of result asa increasing hasbeen acidification , or lowering or , (the amount of heatre (the amount of of the ocean can of pH biogenous , making , Global ------per 3000 m of the world ocean during the latter half of the of oceanduring the latterhalf the world mof 3000 per in the up 0.037°C of increase culated ameantemperature Scientists have cal the atmosphere. heatfrom amount of by andstoring an immense theatmosphere absorbing of the warming carbondioxide, it also has been moderating of by huge effect quantities absorbing greenhouse increasing the While the ocean has been moderating the Earth. ance of moreover, makesit the dominant component in the heat bal heat, and store toabsorb capacity ocean'senormous The by affected in temperature. changes greatly are organisms tomarinelife,asall importance crucial is of ments. This environ to terrestrial in marine environments,compared stable and homogeneous relatively remain temperatures in the ocean, means that water volume of the tremendous coupledwith in temperature, increase heat without a large amount of alarge toabsorb water of ability byThe 1°C). effect is intensified byeffect thehigher the poles (Fig. 5).This andless toward equator and tropics, at the er, more and sea surface the atmosphere warming is great surface) striking the Earth's solarenergy amount of so focused ontoasmallerarea, are its rays overheadat the equatorthan at higher latitudes, directly more tolocation.Becausethesun is according varies temperature Temperature in complex ways in the ocean. Surface varies inhabitants. globally, Earth's climate all of thus affecting and weather patterns regulates in turn the ocean,which of in the circulation changes produce Changing temperatures oxygen for effects, example decreasing indirect solubility. also has many in manyits inhabitants ways. Oceanwarming themarineenvironment and affects directly temperatures excess heat byWe theocean. ocean will thatchanging see of has limits andconsequences, sotoodoesthe absorption excess carbon dioxide by the ocean of Just as the absorption to theatmosphere! transferred were in ocean temperature for this increase all the heat required if global atmosphere, the of 100°C of increase would equal ameantemperature ocean the world of 0.1°C of increase a mean temperature To scientists calculate that warming. put this in perspective, global global system during this period of ed tothe entire all the excess heat add of 84% in ocean, fact the world by absorbed heatenergy amountof anincredible reflects twentieth century. may seemsmall, it Although this increase albedo (reflectivity) of the of (reflectivity) insolation (the ------23 Lesson 3: Pelagic Environment - - - - - per (C&H Fig. 3.22). At Fig. (C&H seasonal thermo (rate of changedepth) with . Thisthermocline a persisis (C&H Fig.3.22a). At polar latitudes, polar At latitudes, (C&H Fig.3.22a). gradient ), are described in detail in your textbook. ), are textbook. your described in detail in temperature profile thermocline tides and may develop within the surface layer of developthe surface within may layer temperate and waves manent thermocline thermoclineso the muchis surfacethe watermuch is colder, 3.22b). In pronouncedless (C&H Fig. at lower latitudes than a shallower however, months, summer cline polar waters,warmed is layer as this increased insolation by tems that produce that tems and other sea-surface them, phenomena ( temperature that have seen We variesnear ways complex in surfacethe of2-dimensional the just is this ocean, the but understand fully the importantpicture. To role temper that aturethe ocean, look below the surface. in we must plays temperatureMeasuring at differentdepths waterthe in col producesumn a temperate and tropical a relatively latitudes, band shallow of warm surface water floats a muchover broader band of surface between the In and deep near-freezing deep water. waters, from about 200 m to about1000 m (660 to 3,300 ft) below the surface, therean intermediate is band of water where the temperature called a steep, is featuretent regionsthese in referredis and the as to ------surface cur (C&H Fig. 3.10). (C&H Fig. Figure 5. Insolation (greater at the equator than the poles). at the equator Insolation (greater 5. Figure El Nino-Southern Oscillation sea-surface temperatures events, which we will learn more about later. Chang events, whichlearn we will more about later.

, which carry warm water from the equator to higher ways, due to the intensifying greenhouse due to the intensifying ways, effect. The conse ofquences changesand ocean these to ocean environments life, and to Earth's climate areand weather systems, of great concern.The major surface currents,sys the global wind eLearning UAF Bouffard, Christen by Illustration (ENSO) the patternses in of sea-surface temperatures (such as with event)an ENSO have dramatic marine life for consequences and produceweather aroundin major shifts world.the Sea- surface temperaturesare unpredictable in changing, now patterns Theseare that systems. global wind sur drivenby face-current insolation. with patterns also vary seasonally, termLonger variations patternsthese in most well, as occur periodic due to notably Because both insolation and the extent of extent and the Because both insolation cover ice and snow sea-surface temperaturesvary vary also seasonally, season Thispicture made is more by complex ally. rents and cold waterlatitudes, the oppositein direction, complex in ice and snow covering the poles, as they reflect as they the poles, covering ice and snow more of the do the darker-coloredthan (and heat) light sun's sea and land imagery Satellite surfaceslatitudes. at lower givesa global us picture of 24 BIOL F150: Introduction to marine biology intermediate layerintermediate on the ocean: a structure three-layered in water sinks,this results (C&H Fig. Because denserwater 3.22a). nent thermocline theperma density mirroring increasing with a pycnocline of for these regions, profiles temperature imagesof like mirror the ocean, at low and middle latitudes, tend to look of much calleda density gradient asteep produces halocline or athermocline of the presence salinity, orwith increasing temperature with decreasing so Density Density: Stratification Stability and the ocean. of sity, structure the vertical anddensity controls its den because they determine importance crucial of are seawater of the salinity and temperature marine organisms, on effects In addition to their direct thermocline. permanent although usually3.22a), orconsistent as the not as steep a called andagradient with depth, sis.Salinity also may vary heats up onadaily ba water shallow depths, asthe surface (C&H Fig.3.22c). abling scientists to identify them and track their movements. abling scientists to identify them and track have unique ocean theworld of regions different masses from water andbottom deep less or fixed, more andsalinity are ture and salinity. temperature that influence Since their tempera tom water masses, two water the ocean's volume. It actually includes comprising of 80% ft), m (5,000 1500 sitting below about dense water cold, of layer deep consists the ocean'svolume. The of 18% about layer, into the layerdeep the surface grades andrepresents pycnocline or where zone, layerintermediate is atransition andsalinity take place.The in temperature that changes here Itis in weather andclimate. seasonal andannual fluctuations by and is affected daily, currents, by andsurface wind, waves layerThis alsocanbemixed exchange with the atmosphere. and gas rivers, of the outflow freezing, tion, precipitation, andlit by the sun, subject to evapora volume. It is warmed the ocean's of 2% about represents ft thick) m or330–660 and a halocline deep layer (the amount of mass per unit volume) increases unit massper volume) increases (the amount of , both essentially out of reach of the processes theprocesses of reach both essentially, outof is seen in some temperature-salinity "signatures" temperature-salinity Diurnal thermoclines Diurnal . The thin surface layer thin surface 100–200 (about . The deep water (also called the pycnocline stratification salinity profiles salinity and the even deeper deeper andtheeven . Density profiles Density pycnocline layer surface layer surface also occur at very alsooccuratvery , which imposesa which , (C&H Fig. bot en , an for ), ------the pycnocline layer, oxygenon theotherhand,while is not In populated. layerisms, becausethis vast is so sparsely light, but by it organ also is not usedupquickly respiring of oxygen by is not produced photosynthesis,due tothelack tion water,by In deep photosynthetic marine organisms. exchange with the air, gas because of andoxygen produc and low latitudes. high near the surface Oxygen are levels oxygencolumn at middle distribution in the water of tern p. 15.28, in a distinct pat also results Stratification 352). (C&HFig. water in deep andhigh levels near the surface with low levels similar to density profiles, ocean oftenappear layer.in the surface Thus, organisms of reach out of bottom sediments, remaining and water essential ocean, nutrients accumulate in deep consequences. Becausemattersinks in the has important other, each from separated effectively which layers ferent a creates Stratification sides of the Atlantic and Pacific Oceans, where winds and theAtlantic andPacific Oceans,where sides of eastern down.The water surface forcing the shore, against is driven water p.15.30, when Downwelling 354). also occurs by called shore coastal winds,upwelling (C&H Fig. a process off driven water surface rises toreplace water when occurs during the spring.use byUpwelling also organisms surface to produce weatherduring the winter can combine andstormy peratures coldtem where regions, in temperate a seasonal process as also is important extremely This up to the surface. back water have accumulated in deep as it brings nutrients which is critical to marine life, which to asoverturn, referred are bymass. thesinking water Together ward thesemovements upwelling below, water causing it to sink. An opposite movement, called densethan the to anextent surface), that more it becomes iceonthe of theformation or evaporation (through creases its salinity or in decreases water surface of the temperature called process dense and sinks, cools, a it becomes more water surface in a relatively tion, resulting pycnocline a strong and stratifica of vents the formation pre generally water surface of the chilling In polar regions, zone in an results which organisms, respiring bybeing generated photosynthesis, by it is being depleted at these intermediate depths (C&H Fig. depths p.16.18, attheseintermediate 372). , is produced as deeper water is displacedup water as deeper is produced , overturn downwelling and mixing, replenishing nutrients for andmixing, replenishing stable water column stable water nutrient profiles unstable water column unstable water . Downwelling occurs whenever whenever Downwelling . occurs oxygen minimum , with , the dif in the open . As ------25 Lesson 3: Pelagic Environment

------in Fig. Fig. in , exerted light profile Pressure pressure profile , and how this varies, and how this with light How seawater transmits important.depth,vitally is also providesSunlight energythe directly and photosynthesis or indirectly photosynthesis, for supportsvirtually readilyis life. Light all transmittedthrough the atmosphere,the ocean reflects but much of the light surface,its striking and quickly absorbsdoes that the light penetrate, converting 7). A it to heat (Fig. aboutthat 3.11) shows 65% ofFig. (C&H abis light visible 1 atm of pressure,pressurethe plus ofofcolumn the water above The them. pressure exerted watervery adds up by about each 1 atm for 10 m (33 ft) ofquickly, depth, since watermuch is (the air heavier than relationship).linear this 6 shows The pressure Chal the in Deep oflenger sea level,below 11 km at Mariana Trench, the thereforewould be about1100 atm, or 16,170 lb per square nearly is inch! Water gases incompressible, are but highly compressible.a cabinat 10 A plane flying km with pressure of 1 be atm must engineered to prevent while it from exploding, same cabinthe with a submarine pressure at depthbe must marine organismswith so as not to implode. Similarly, built gas-filledstructures and air bladderslungs like con must extremetend with pressure fromwhen without, at depth, and extreme pressure fromwhen within, waterthe in rising col umn. Sperm whales, which can dive down more2000 than m backand surfacethe to may up space ofthe within hour, an extrememost to the be subjected pressure changes of all. veyor is vitally important vitally is veyor to carries life, as it to marine oxygen deep watersreturnsand surface.the to nutrients has also It influenceimmense global weather patterns on and climate, transportsit as fromheat tropics the toward and poles the vice versa. Theof possibility thermohato the disturbances circulationline of resulta as ocean, the of increasingthe greenhouseeffect and global warming,of is great concern. and Sound Light Pressure, Depthwaterthe in affectsalso column several other impor properties physical tant of seawater. of weight the by pressingcolumn the fromdown above, in hardlyofweight the notice creasesdepth. with We steadily enormousthe of column exerting air above us, 1 atm (atmo sphereper square1 kg = cm, or 14.7 per squarelb of in) pressuresea level. at Marine organismsare this to subject . ------ther (C&H Fig. (C&H Fig. that circulatethat wa is formedis streams and NorthDeep Atlantic great ocean conveyor ocean great (C&H Figs. 15.33, p. 357), 15.33, p. (C&H Figs. coastal upwelling coastal . It beginsof the polar seas south in subsurfacecurrents Antarctic Bottom Water equatorial upwelling equatorial streamsSouthern to the way the all south, Ocean or all the rain fallingon the planet. The great ocean con veyor belt, this circuit belt, this the called is veyor perhaps of thousands Requiring years ofthe completion for one cycle,scale. Thea global ocean on the mixes it volume of calculated to equal 100 is waterAmazon Rivers, involved Water 3.24). ThisdeepFig. (C&H circulationshal with connected is lower circulation, returningwarm surface-water from the Oceans to the North and Indian Pacific Ocean, thus Atlantic 3.25). con an immense Like a circuitcompleting (C&H Fig. formsentrywaythe a dam at as acts and a sill a deeperto basin. The dense water overflowsit eventually piles up until streamsand sill the deeper the into and down out In basin. way, this Similarly, north,equator. past the Greenland and around Antarctica, relatively in basins. shallow Here, surface and water becomes very cold and very salty, therefore veryhapthis how see will (we sinks dense, so it pens whensea ice, below). This we discuss watersinking becomes trapped of edge at the basin, whichshallow the slowly, in immense immense in slowly, ter throughoutworldthe deep-ocean this ocean. Because differences density circulationdrivenis by tem produced by peraturechangessurface, and salinity at the called is it mohaline circulation upwelling and downwelling, areand downwelling, upwelling important arethey not but formthe dominant of circulationthe ocean. their in Using temperature-salinity"signatures," have tracked scientists deep- and bottom-water and discoveredmasses that these water masses (80% ofare the ocean's volume) very flowing, Density controlsDensity the verticalonly not structure of ocean, the circulationthe also ofbut vastthe of majority volume. its The types of waterso far, we have discussed movement surface currentsand the vertical flow of in water involved surface currentspile up surface waters, The sinking. inducing whereformeris it process occurs at the equator, primarily called and the latter often occurs at about 30°N and 30°S latitudes. surface currents surfacetend to move water fromaway the coast, are importantareas of occurs upwelling when water rises 355). Similarly, 15.31, p. to replace surface waterdrawn apart diverging two by sur face currents,occurs and downwelling where converging two 26 BIOL F150: Introduction to marine biology Figure 6. Water pressure increases steadily withdepth. increase. This produces a region of minimum sound velocity, of aregion produces This increase. continues to as pressure depths, at greater then an increase thermocline, thepermanent of in the region decreases ature column, as temper in the water in velocity with depth crease ingly, salinity,Accord with increasing andpressure. temperature sound increases speed of lesson. The in the previous learned the ocean, as we and humans employ toexplore sound waves sonar), sound (nature's for communication and echolocation use "noise pollution." Marine organisms human-generated increasingly, and, by sounds produced marinecreatures with with is alive sea m/s). The 340 m/s) than in air (about 1500 Unlike light, light (C&H Fig. in 3.12). natural objects at depth, of pearance and theblueishap thesea, greyish or of colors and green blue light. This than readily more absorbed are thespectrum, end of red Longer atthe with wavelengths, light depth. also changes of quality The highest near the surface. are densities generally and organism the ocean, of place only in the upper reaches photosynthesis takes As a result, m. to100 penetrates 1% water, in clear the surface and less than within sorbed 1 mof sound velocity profiles sound velocity sound selective absorption selective travels faster and farther in water (about (about in water andfarther faster travels (Fig.8) show an initial de accounts forthe blue - - - - Figure 7. Lightisquickly absorbedasitpenetrates water. Icebergs, Ice Shelves Ice and Islands in this zone. mi), have beenrecorded 15,500 km (about as 25,000 oceanbasins,as far entire across transmissions and man-made sounds. Sound vocalizations, earthquakes, m down, at low 1200 and middle latitudes), to monitor whale the SOFAR (600– channel equipment atthe center of cording atunnel. Scientists place re through echoing as if Ranging), this layer, calledthe in energy distances without losing much great can travel the layer. the centerof focused, sound waves toward Thus andbelow, above (bent) from directed andare refracted are becausethey here "trapped" minimum sound velocity are of oxygen in this layertraveling minimum Sound waves zone. velocity, greater like the much of betweenlayers sandwiched forms at the surface, insulating and maintaining the liquid atthe surface, forms for marineandaquatic life, since ice is crucial floats. This ter becomes less dense as a solid than as a liquid, so ice remarkably, Even more wa its unique properties. another of the planet in all three Water's ice at the surface. to coexist ability on of presence environment is the the pelagic of feature Another important SOFARchannel phases —gas, liquid —gas, and solid—is

(for SOund Fixing And Illustrations by Christen Bouffard, UAF eLearning UAF Bouffard, Christen by Illustrations - - 27 Lesson 3: Pelagic Environment ------, at Ant Terror and Ross Ice Shelf Ross Erebus ), aboutofsize the France.was It 2 in 1912, in off ofcoast the Newfound Titanic areattached found to 44% ofAntarctic the (182,000 mi 2 Their ice covers areas, immense is 100–1000 m thick, and ofthousands years old. The largestthe is 472,000 km discovered1841, in the during arctichonor of in and later named expedition, commander. its response(Ross' was, on seeing it "there's no more chance of through sailing throughthan that Cliffs ") Ice the of Dover. open water, icebergs are transportedopen water, currents, by primarily large due to their underwater relativelyand extension small "sail" area. They become lockedthe sea win ice during into and are released continually they Since againspring. in ter, large a relativelyicebergs only few, melt and erode away, to be carried enough long last whichlanes, shipping into 2–3 take years. may was It of one persistentthese bergs the sank that North the been has Atlantic disaster, that since Ever land. icebergs are monitored patrolled to locate icebergs. Today, worldwide,remote using sensors on polar orbiting satellites. Ice shelves coasts ofand the coastline, Greenland and northern Elles mere Canada. Island, They areseaward the created by ex ofpansion glaciers,accumula snow annual augmented by ofa tongue until tion, sea surface. the over out ice extends - - - - Figure 8. The SOFAR channel. The SOFAR 8. Figure

, or just "bergs,", or just calve (break off) from glaciers danger to vessels. Large bergs to vessels. be surroundeddanger may fog by differencethe due to banks, temperature in ice between the run They may surroundingand the aground, and can water. preventingthe installation the sea floor, scour and gouge of underseasome areas.in and pipelines cables Afloat in eLearning UAF Bouffard, Christen by Illustration Since 60–80% Since of this a berg'ssubmerged,is however, bulk literallyis oftip the just iceberg! the Icebergshun can weigh dreds ofof thousands and are tons, unstable, dangerously to breaktending or capsize. Subsurface "rams" projecting part visible the far beyond of berg the aregreat another sea surface by 5 m acrosssea surface surface) the sea by and "bergy bits" 5–14 to "verym) (1–4 m by large over bergs" 75 by m (over 213 m). Theicebergknown tallest the North in tow Atlantic ered 168 m above the sea, ofthe height a 55-story building. Icebergs whichcover sheets ice huge the flow sea, primarily to the Antarcticaing and Greenland. shapes They many come in rangingand sizes, from "growlers"1 m abovethan (less the environment below, and it melts quicklymelts when it and temperatures below, environment rise again. gen two in environments marine in is found Ice eralforms. Freshwaterglaciersin land on originates ice and the ocean as icebergs,ends up in and ice islands. shelves ice The second form,ice, occurs sea whenseawater freezes. 28 BIOL F150: Introduction to marine biology ner, beneath ( being melted from as they are thin become And iceshelves are common. becoming more calving events, alsois normal from a phenomenon different shelves, of disintegration extent. The in ice shelf al decrease agener the twentieth century, example of themostdramatic byduring reduced 90% Island iceshelves were all, Ellesmere Island In in 2008. Ellesmere of off one, alsocalvedentirely took lessthan an hour. IceShelf, Markham another huge The Manhatten.Caughtby satellite imagery, this amazing event of becoming an ice island 2005, the size 13, Island on August Ellesmere of off fractured Ayles Ice Shelf entire The 1978. station with plant complete and runway,power 1952- from amanned research Island, carried Ellesmere of calved off islands called ice shelves are that calve from Huge,icebergs flat in this region. productivity in a loss of system and resulting byin 1841 theRossered expedition), upsetting the eco the Ross Sea (also discov seaice out of of flow normal the dams that blocked formed this huge berg piecesof The calving. within months of apart breaking it began before km 11,000 It was 2000. of in March theRoss IceShelf off broke B15, Iceberg recorded, ever icebergs the largest Oneof shelves also calve icebergs. a volume of water equal to the volume of ice above the iceabove equaltothe volume of water a volume of displaces ice already becausefloating level, sea raise rectly andice shelves does not di icebergs creasing. Melting of to thesea, runs land glaciers from expand,and meltwater waters surface poles.As warming a exposedwater. by ing absorbed, thedarker produces This heatbe and more in less light being reflected, results age to thehigh fluencing beneaththem,thus in thewater and salinity of perature thetem regulate They organisms. unique communities of Ice shelves support ships and oil platforms. and hazarding marine lifelocally,consequences,in addition to disrupting iceshelves may haveglobal of and disintegration Thinning change. and climate global warming effect, ing greenhouse linked to the accelerat are these changes to the poles. All of water bringing warmer winds and currents and tochanging waters, and Antarctic Arctic of warming due tothe gradual positive feedback positive . Some have even been inhabited. T-3, beeninhabited. . Somehaveeven an iceisland thermohaline circulation thermohaline albedo of polar areas. Decreasing icecover Decreasing polarareas. of effect, accelerating warming at the warming accelerating effect, 2, about the size of Connecticut, thesizeof about sea level . They alsocontribute. They basal melting alsois in ice ), ------surface of a calm lake. The presence of salt in water lowers lowers saltin water of presence The calm lake. a of surface onthe In this manner, icewill form athin, glass-like sheet of at0°C. occurs andfreezing andthen density decreases 4°C, tothe point of its density increases both locally andglobally.important cools, water When fresh more icefound inandfar the sea, accounting forof 95% seawater, from complexforms andextensive, more andis far Seaice different. but glacialice, itactually from able isvery To eye, theuntrained Ice Sea effect. greenhouse duetotherunaway changes celerate a the ocean,anothercaseof toward will move faster held back the glacial icethat was off,When they break ice shelves act like dams onglaciers. water.However, since sea level, increase it can indirectly ice, or summer.to someextent each landfast includes ice pack The melt. The of tudes,the winter. each again spring andforms melts each At higher lati winter icepack continues. In this way,as freezing the ice fields andeventually massive ice floes larger unite to form pieces,it called "pancake" or"pan"ice,which into circular andshape brittle. Wind action break and wave becomes more to the bottom(oppositetohow glacial ice accumulates), and ice by thickens theadditionof crust similar thickness.This of ice unlike man, a freshwater gull a but notof the weight of and wind, and is able to support to waves bends in response called "nilas,"which cm thick, ice5–10 new of flexible crust a creates freezing Further during this process). that occurs ice"is one form ("grease oroily acloudy appearance surface thesea gives congealing slush, which layer thick of paratively in com a floating sinks as itcools,with surface ice crystals atthe water As a result, at–2°C. as it cools, until it freezes continues to get denser soseawater –2°C, incide at about point co maximum density and the freezing of temperature (the freezing maximum density and of bothof the temperature multiyear ice freezing point freezing fast ice fast summer ice pack summer permanent polaricepack permanent , which is attached to the shore, and tothe shore, is attached , which ) is formed, consisting of consisting of ) is formed, , which has survived at least two summers' at leasttwosummers' hassurvived which , ). In seawater of average salinity, average the of ). In seawater sea ice sea also remains but is broken up but is broken also remains feedback loop mayindistinguish appear maximumdensity seasonal icepack occurs, consisting occurs, annual ice that will ac drift ice that (or , at - - - - - . 29 Lesson 3: Pelagic Environment ------due to shrinking ice due to shrinking . Sea ice reflects 80% of albedo feedback loop feedback predicting ice-free be completely Arcticthe that sum in may first the for 2030s, the aboutmer by in time years. a million are poles the that know warmingWe faster restthe than of One we are planet, but the trying still to understand why. logicalreason decreasingthat is is creatingcover a whileit, hits that sunlight the open water absorbs 90%. Huge global heat balance. Alarming changes to the Arctic ice pack and partsof Antarctic the pack,ice changesto the similar been observedhave ice shelves, seen with recent in de freezing it’s thinning, is Sea ice later and breaking cades. and therebeen a dramatic has declineup earlier, ex the in oftent coverage ice Arcticthe in partsand of Antarctic.the the Arctic ice in Multiyear shrank an average of 8 percent per begandecade, monitoring satellite since the late 1970s, in the beginninguntil of whenbeganit dropping century, this September In ofmore precipitously. 2007, a record minimum was extent previousthe seen, 40% than less aver 28-year age. (Thearea smallest the is extent minimum ice covered by Thisre summer melt season, during the March-September.) cord was brokenSeptember in 2012, whenex minimum the oftent Arcticwassea ice nearlyhistorical the than 50% less average.words,other In Arcticaccel at an melting is sea ice eratingrate, fastermodels had predicted,most scientific than some to a "death spiral." by models are and likened Revised blocks of ice areand currents, wind driven by such with speed and energyoverride violently they that shore the or landfastgroupice. A of hunters fast the on office Barrow, abandon to had Alaska, runand dogtheir gear and teams ofthe path in lives, their for an ivu1957. in Hundreds of was a family crushedyearsivu,an earlier, by home their in remainingbodies their frozen Arcticthe in a reuntil soil cent storm unearthed on the coast near Barrow. the site, of one This surprisingly, challenging is, environment eco the richestlogically and most importantthe world. in Sea ice is and diversehome to unique of communities on marine life, which other organisms many (including depend,humans) as course.this see later in we will Thepack—itsice extent, oftiming and the movements, its freeze-up and melt—is but regionally, conditions physical and life to only not critical because ofrole its circulationocean in globally, also and . ------ther brine rejec , long cracks, long in , piles of, piles jumbled Leads great ocean conveyor ocean great near the near poles that drivethe pressure ridges " by Inupiat Eskimos, occursEskimos, Inupiat " by when and the , especially when covered with snow, when, especially covered snow, with ivu , occur over upwellings of upwellings , occur over relatively warm albedo polynyas , this process, this resultsfreshin ice near the top of the deep-water masses to as push ice or " ice push to as tend even farther downward from the bottom of the pack. Thesesea floor and plow the shallow in can gouge keels areas. Beachesarealso ofactions the shaped by sea ice. One ofimpressivemost the sea-ice phenomena, referred called leewardthe on or of side water, whereand coasts, islands prevailing fromice away the keep winds blowing shore. Col ice floesliding up push blocks of ice 10 m or more ex may "keels" that with high, nate growth and melting, and to movement and deformationand movement to and growthnate melting, and and currents.tides wind, by Ice floes are carried thousands of kilometers from away ice pack. the passage ofallow to enough wide ice the or a whale,a ship open and close back up again. Other areas of open water, blockingaccess to the water's heat stores. Because huge ofhigh its sea ice also reflects from heat away the Earth'ssurface. and variable, extremely complex is Sea ice to alter subject mohaline circulation Sea ice is important to regional and global heat balance as well. Theother ways, in ice pack heat exchange inhibits sea fromthe insulates It the air. sea and the between the makes the air colder by and actually a blanket, like cold air, tion pack and increasinglythe lower layers.ice in salty the As cold water below the forming saltier it also gets ice gets to denser and beginsbecoming a major contributor to sink, the As seawaterAs freezes,be incorporated can salt the all not expelled the ice, into and extremely salty liquid—brine—is waterthe into formingthe beneath ice. Called to the bottom during each winter. Thus, the youngest lay Thus, the youngest to the bottom eachduring winter. ers arecontrastbottom, at the in glaciers, to mi and they grate upward time. The over migration of each new layer take about 12 years, top may to the contrast also in to ice presentglaciers, in whichof be thousands may years old. Newa thickness ice attains of 30 cm to 2–3 its m during 10 than m less usually ice is multiyear Even first winter. thick, muchglaciersthan thinner because and ice shelves, undergoesit cycle a continual ofand replacement. loss The whilenew added ice is melts eachsurface summer, layer 30 BIOL F150: Introduction to marine biology the ocean. Water masses are in contact along "ocean fronts" the ocean. Water in contact along "ocean fronts" masses are "weather"in them as examples of coming to view ments are study thesemove who plex ways.Physical oceanographers com andextremely this 3-dimensional environment in varied moving in masses are water being static, from seen that, far linity,density, temperature, andlight. We alsohave pressure composition, sa chemical including to many characteristics, with respect differentiated stead is horizontally andvertically We homogenous, but from in haveseenthat the ocean is far WeatherOcean andglobally. conditions regionally changing as adriver, of bothasanindicator, seaiceconditions serve Thus, and worldwide. may weather systems and climate also influence like the Moreover, these regions. disturbances to currents through impacts on conditions and lifein sea icewill have enormous theloss of the mechanisms, factor. of important Regardless may more an even be tothepoles,which energy porting trans are weather patterns ies alsosuggest that changing Recent stud and melting at the poles, andtoglobalwarming. warming may contribute both to accelerated mum) therefore sunlight is at amaxi losses in icecoverin summer (when passes over the Rio Grande Rise, off the coast of Brazil. thecoastof off Rise, passes overGrande the Rio e.g. submarine waterfalls, as it numerous tom Waterforms conveyor,ocean in the great On its way north Bot Antarctic found onland. any waterfall can dwarf produced cataracts Water, Atlantic Deep Water andNorth the discussed above), Bottom Antarctic of onto the other side(as in the formation in the sea floor,lates behind a barrier eventually spilling over sinks and accumu water When dense surface this effect. of aspectacular example are mountains. Submarine "waterfalls" like by affected landforms same ways that air masses are masses, in the water of the flow alsoaffects topography floor density. greater water's Sea periods in the ocean, becauseof slowly andoverlonger more time but operates atmosphere, Turbulence asit is in the alongoceanfronts, is generated mass. another water against Stream like the Gulf current face asur the abutment of or flood, a flash after ariver mouth of the from a mud plume spreading is the edge of ocean front an An example of in our atmosphere. similar to weatherfronts Gulf Stream Gulf and to thermohaline circulation thermohaline , it ------the surface down to about 100 or 200 m, including nearly all nearly m, including or 200 100 downtoabout the surface m deep.1000 shallowest zone is the The and the deep; into two zones: the is divided tively.the bathyal Over environment benthic the pelagic zone, m deep, 4000–6000 zones,and from m, below 6000 respec abyssopelagic deepest The to depth. according divided further zonesare These break. beyond open water theshelf tinental shelf, andthe the regions, general into two isdivided realm environment (Fig. pelagic The 9). basin (Fig. zones found 3)with the habitat in the pelagic seawater, tofill in our benthic ready weare acteristics of Now that wehavedescribedthe physical char and chemical Pelagic Habitat Zones cinating life forms they support, in Part 3 of this course. this course. of 3 in Part they support, cinating life forms vents). We these zones, will and thefas explore thermal hydro heat, around andcold(orextreme pressure treme nosunlight at all and is subject to ex receives realm vast This zone." sometimes elided into a single "deep-sea are which abyssopelagic andhadopelagic, ing thebathypelagic, Below the dysphotic lies the sure. cold andpres subjected to increasing are and organisms photosynthesis, but not enough to support light penetrates some is the "twilight zone" where This to themesopelagic. photic lies the nutrients. sufficient Below the given thesis. here, Lifethrives photosyn andsupports waters thesurface sunlight warms The dysphotic zones. into photic, and aphotic light penetration, to according environmentclassified alsocanbe pelagic The the open ocean. of waters the neritic zoneplus the surface photic zone mesopelagic zoneslie over thehadalandabyssal benthic dysphotic zone dysphotic corresponds to the epipelagic, where where tothe epipelagic, corresponds neritic zone oceanic zone , from about 100 or 200 m, to m, 200 or 100 about from , bathypelagic , roughly corresponding corresponding roughly , , which lies overthe con which , aphotic zone , which includes all the includes , which hadopelagic epipelagic , 1000–4000 m , 1000–4000 , includ , from , from and ------

Epipelagic Mesopelagic Bathypelagic Abyssopelagic Deep Sea Deep Pelagic

Oceanic

1000 m 1000 4000 m 4000 100-200 m 100-200 Hadopelagic Neritic Hadal Abyssal Bathyal Subtidal Shelf Break Intertidal High Tide High Tide Mark Benthic and Pelagic Habitat Zones Habitat Benthic and Pelagic Low Tide Tide Low Mark Benthic

31 Lesson 3: Pelagic Environment

Epipelagic Mesopelagic Bathypelagic Abyssopelagic Deep Sea Deep Pelagic

Oceanic

1000 m 1000 4000 m 4000 100-200 m 100-200 Hadopelagic Neritic Figure 9. Benthic and pelagic habitat zones habitat Benthic and pelagic 9. Figure Hadal Abyssal Bathyal Subtidal Shelf Break Intertidal High Tide High Tide Mark

Low Tide Tide Low Mark Illustration by Christen Bouffard, UAF eLearning UAF Bouffard, Christen by Illustration Benthic 32 BIOL F150: Introduction to marine biology your textbook. of attheback theglossary guideandinyour textbook, andcheck thematerialinthiscourse re-read theseterms, any of themeaningof about unclear you're If for understanding. for thislesson.Doaself-check somekey terms Listed beloware • • • • • • • • • • • • • • • • • • • • • conveyor andgreat ocean circulation thermohaline masses andbottom water deep water downwelling, upwelling, andoverturn stratification cline andpycnocline halo thermocline, permanent thermocline, etc.) pressure, temperature, (of profile insolation andalbedo heat capacity pH andoceanacidification global warming and effect, greenhouse gas, greenhouse exchangegas Niskin bottle calcium carbonate pollutant nutrient constantproportions of rule salinity (psu,ppt) salt andion solvent universal andsolute solvent cycle hydrologic glossary terms - • • • • • • • • • • • • • • • • • • • Coriolis effect andEkmantransport Coriolis effect ridge andivu pressure lead, polynya, ice drift iceandfast annual iceandmultiyear ice icepack and permanent winter/seasonal icepack,summerpack andiceisland iceshelf iceberg, point freezing SOFAR channel oxygen minimum zone photic zone, dysphotic zoneandaphotic photic zone, dysphotic zone lagic zone, zoneandhadope lagic abyssopelagic zone, zone,epipelagic mesopelagic bathype neritic zoneandoceanic tides tidesandsemidiurnal diurnal tsunami seas, swells andsurf fetch gyres current Stream andGulf equatorial currents andpolareasterlies winds,westerlies trade - - 33 Lesson 3: Pelagic Environment - in your in your . For example, if example, . For to asked complete C). Describe 5 other important differ o and clear, but also but clear, and • brief out of retirement Undersea at the National (she's now on display Trieste C) is lower than that of freshwater ice (0 o (Please make your answers and concise) complete your make (Please Written Assignment LessonWritten 3 for and submit the written assignment to UAF eLearning to UAF assignment written the and submit according Education & Distance to directions found , not those ofthose , not or Coursetextbook the Be Guide. cover sheet pack in recent decades. Speculate on how these changes could affect the great ocean conveyor, and how altering the pack in recent changes Speculate on how these could affect decades. the great ocean conveyor, as if to be very (Try could have global consequences. great clear, to someone who explaining ocean conveyor has no ofknowledge terms the are understanding in order you to show your using, of processes.) these complex The freezing point of sea ice (-2 ences between sea ice and glacial ice. Explain the role of brine rejection, which occurs as sea ice freezes, in the formation of deep-water and how masses, Describe some specific changes that have been observed in the Arctic ice it helps to drive the great ocean conveyor. warmed it from here, (step- the sun, rising and being displaced, which continue explaining by creates Taking winds. how the Gulfby-step) Stream current develops carries and warm water from to Norway. the equator all the way Include effect, the Coriolis Ekman transport and gyres explanation. in your semidiurnalMost coasts experience what Define these tides and explain causes them to occur. tides. at 50 m, 500 m, 1000 m, 5000 m and 10,000 m deep. at 50 m, 500 m, 1000 m, 5000 m and 10,000 Off the east coast of Greenland, pack ice covers while most of at about the sea for the same latitude and the year, Norwegian ports a fewonly hundred remain miles away, ice-free and temperatures This are is considerably warmer. due in part to relatively warm waters from the Gulf Stream current coast, while bathing Norway's the East Greenland current brings frigid water from the Arctic Ocean to Greenland. The Gulf Stream begins with air at the equator being Imagine that you take the bathyscaphe take the bathyscaphe Imagine that you Museum in Keyport Deep of trip down to the Challenger another for Washington), tropical in the the Mariana Trench, waters of the western (or diagram) Describe Pacific. salinity and light conditions the temperature, pressure, density, observethat you measurements in the water at the surface, and surrounding descend. Take submarine as you your Complete the crossword All the terms page. puzzle on the following in the puzzle are included in the "Glossary (Submit the completed puzzle, or a list of this lesson. for Terms" the answers, with this written assignment.) in seawater there?How do the ions found get (Describe 4 ways.) 6. 7. 5. 4. 3. 1. 2. online at eLearning.uaf.edu and in the course received.online at eLearning.uaf.edu materials you own words understanding, be clearyour and show are imagineto you that explaining (To list. just do so - don't "describe" something, someone who of has no knowledge the subject.) Includea Study the information presented for this lesson until you feel that you have a good understanding. Write your answers your information the understanding. have a good Study Write feel that you presented you until lesson this for 34 BIOL F150: Introduction to marine biology 35 Lesson 3: Pelagic Environment (Tear out and submit for grading) out and (Tear crossword puzzle for Lesson 3 for puzzle crossword 36 BIOL F150: Introduction to marine biology 37 Lesson 4: Biological Principles, Cells, Organisms & Ecosystems - . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). be found at the publisher's Online Learning at the publisher's be found Center: www.mhhe.com/castrohuber9e Know the basic characteristics of in which the fundamental ways living things, they these attributes. basis for and the molecular differ from nonliving matter, Be familiar with the general structure of types of and the basic ecosystems, interac tions among organisms and between organisms and their environment. the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education. 3. 4. 5. 211–229) 64–83) and Chapter 10 (pp. C&H: Chapter 4 (pp. More information, including interactive flashcards, quizzes, videoclips can and web-links, 1. 2. Reading Assignment Reading Objectives for this Lesson this for Objectives Lesson the Completing Steps for Marine organisms are diverse incredibly in form and function, with diverse strategies for coping with the challenges their surroundings. presentedmeeting their needs and for by They interactions, also are in complex organisms both with other involved and with their are basic characteristics, however, Underlying environments. complexity, all this physical needs and challenges all organisms, shared which and general by principles by organisms interact In this lesson, we review with each other and with their environments. these fundamentals of biology and ecology. Principles, Cells, & Organisms Ecosystems Biological Biological 4 38 BIOL F150: Introduction to marine biology N elements capable of forming strong bonds with each other, bonds with each strong forming of elements capable tobe thelightest four happen These and nitrogen. carbon hydrogen, oxygen, matter is made of living of 99% crust, the most abundant elements in the Earth's are num and iron oxygen, life.Whereas silicon, alumi of essential ingredients elements,are only 30 about occurring the 90-plus naturally chemicals. Of of aunique assortment consist of ganisms or life? At the most basic level, the unique attributes of are molecules, andwhat distinguishes these collections of what andphysics.So, chemistry to thesamelaws of conforming ecules—the same molecules found in inanimate matter, downinto simple, lifeless mol thingsbroken canbe Living Diverse, Complex Organized and this course. 3 of systems, and its 2, ecological in Part in Part inhabitants, for meeting the ocean's diverse in preparation andecology, biology these fundamentals of son, wereview other andwith their environments.act with each In this les inter organisms principles by andgeneral which organisms, by shared all needs andchallenges basic characteristics, are all this complexity,complicated—it is! Underlying however, ecology and with their physical environments; with other organisms too, involved both organisms, in complexare interactions, Marine ment, but also with the landandatmosphere. not only complex within the marineenviron interactions, that the oceanis involvedWe in alsolearned surroundings. byneeds and for coping with presented the their challenges for meeting their strategies andfunction, with diverse form in too, incredibly diverse challenging.organisms, Marine are dynamic and that the marine environment is varied, extremely Getting organized: organisms &ecosystems Cells, is the study of these interactions. If all this sounds If theseinteractions. is the study of (living) aspect. In the previous lessons, we learned lessons, welearned aspect. In the previous (living) toits we turn theocean, of ing) features with the familiar Now that we are abiotic (nonliv biotic • ------

sizes and shapes of molecules, including very large, com large, very molecules,including of sizes and shapes of variety bythe greatest far forming of capable and are other andwith other atoms, at bonding with each versatile together. the most all the elements, carbonatoms are Of carbonatomsbonded two or more consisting of "backbone" compounds by their carbon inorganic distinguished from are ways toproduce at combining together in myriad adept supremely and are proteins. Similarly, all DNA created and RNA molecules are different of the endless variety sequences,to create different of linked in together chains amino acids are 20 things.These of bloc smaller, of limited variety avery from constructed found things is in living macromolecules of limitless variety molecules they contain. Yet, organic the virtually the kinds of let alone all life on Earth, fully know, types of all the different acids. Scientists have only to identify, begun andmay never and nucleic proteins thing has its own unique complement of living type of found each in a human. proteins In fact, ferent the approximatelydif 35,000 is identical to anyproteins of these proteins.None of thousand types of several including molecules, organic different bacterium contains thousands of amicroscopic simple molecules.In contrast, of mixtures dom ran usually consisting of organization, has little structural versity its extraordinary life are most obvious qualities of The these molecules and their interactions. of the properties and lipids cules of assortment andavast water matter, organic is of composed which of plex onescalled amino acid amino RNA k molecules. All proteins are formed from just 20 kinds just 20 from formed k molecules. are All proteins belong to one of four major classes: fourclasses: major belongtooneof , and the macromolecular and the macromolecular , , complexity ) and organic molecules organic proteins molecules, which are identical in all living identical in all living are molecules,which organic compounds organic macromolecules , and . All of life's attributes stem from life's attributes stem from . All of organization . Most of these Most of . nucleic acids . Living things consist . Living . Organic compounds Organic . . Inanimate matter carbohydrates building- ( mole DNA di - - - - - ,

39 Lesson 4: Biological Principles, Cells, Organisms & Ecosystems ------are . This polypeptide mitochondria , differenttypes of for tRNA informational macromol . Translation takes place on the place on takes . Translation assemble themselvesassemble , whichdetermined is par its by tRNAs ofnucleotidetheir into sequences translation conformation transcription , in turn, in serve construction the as templates for of . Theofsequences molecules nucleotidesDNA the in , is unstable in its extended form.extended its in unstable , is Eachpolypeptide tion-rich. The particularamino acid sequence of a protein determinesstructure,3-dimensional specific its whichin the wateryIn turn determines specific biologicalits activity. of environment cell, the long strand the of amino acids pro translationthe duced by of a called mRNA, chain curlsspontaneously a into specific 3-dimensional and folds shape, or Nucleicacids and proteins are ecules ofencode the instructions a cell constructingfor oth the all serve molecules DNA components. er cellular as templates the for complementary nucleotidestrandsin sequences of RNA. Some ofor RNAs called messenger molecules, RNA these mRNAs proteins, during whereribosomes, are acids amino spe together in linked correspondingsequences, cific code ofthe to nucleotide strands mRNA the in sequences translated.being Amino acids are carried molecules, other RNA to the ribosomes by called transferor RNAs the instructionseachtype of encoded way, this amino acid. In aremolecules DNA in used to construct protein molecules. proteinOnce assembled, also become molecules informa transmitinstructions the construction the for and functioning of an organism, and proteinsare the instruments carryfor instructions.these out ing Each has component also cellular the example, For a specific role to play. manufacture and ribosomes power plants, cell's the proteins. Each type of(muscleand tissue cell or nerve) serves a spe purpose,cific as does eachorgan (heart organ or kidney), system (respiratory gross or and digestive) structural com ponent (a branch, or arm)an a wing, the organism. within Self-Assembling organized, and highly Complex both structurally and livingarethings furtherdistinguished functionally, capacity their to by abilityunique properunique the in originates also ofties interact.they ways and the biomolecules, , , ------pro cyto to form species , suchas (Fig. 10). (Fig. organ systems , forms of life con organism . Organellesother and cell membrane , respectively. In all, there In , respectively. cells (C&H Fig. 4.8). Mul (C&H Fig. cells . Supramolecular complexes, , various into assemble tissues multicellular organelles molecules, repeating molecules, chains in of fatty acids eukaryotic tissues supramolecularcomplexes ribosomes , and , consisting of, consisting cell. Most are a single and nucleotide , such as bacteria, whosecomponents are cellular glucose , and organs are combined into —which surrounded is a by , the basic unit ofunit basic , the life. The vastofmajority livingthings unicellular cell turalframeworks. The storeand RNA, nucleic DNA and acids, Every structuralofcomponent organism an a specific has also serve Biomolecules function. in the same functions basically Carbohydratesstore and lipids cells. all fuel and form struc sist ofsist more complex formsticellular differentcontain types of cells Similar cells. group together into organs whichindividual together make up the a are karyotes organized not usually organelles into 4.7). More(C&H Fig. and all unicellular, complex membranes turn,in be organized may more into membane- intricate, structuresbound called structures are organizeda watery within matrix—the plasm The diverse,of molecules complex organicmatter are assem even into more organizedbled highly complex, structures in Thislivingand orderliness, things. complexity again, is based Macromolecules underlyingan on and economy. simplicity are into combined shareofsets similar proteins fromnucleicand distinct acids, those of we can surmise other species, these character that ofsets istic macromolecules areof the identity the basis for differentthe forms of diversity. astonishing its for life, and is based on an underlying simplicity and economy. Further, Further, based is on an underlyingand economy. simplicity formsall since of molecules, same building-block the life use since sharethey we can infer that a common origin. Finally, individualsall of a certaintype of organism,or other relatively small, organic compounds that can be found virtuallyin Thesecells. all universal organiccompounds are capable of producingvarietythe limitless of more complex can see, therefore, We organicin found molecules matter. diversitymolecular the that of and complexity livingthings from5 just various Large carbohydrate molecules lipid and sequences. of also consist chainsof sug simple usually blocks, building ars like are around100-200 ofkinds blockand molecules building 40

BIOL F150: Introduction to marine biology

Figure 10. Complex organisms, like this seaweed, self assemble from simple molecules which are organized at many levels. many at organized are which molecules simple from assemble self seaweed, this like organisms, Complex 10. Figure

Cellulose Sugars Cell Wall Cell

Organism

Amino Acids Amino Protein Plasma Membrane Plasma

Organelles and Cells and Organelles Chromosome Nucleotides DNA

Precursors

Molecules Complexes

Macromolecules Block Building Supramolecular Supramolecular 41 Lesson 4: Biological Principles, Cells, Organisms & Ecosystems . , ------molecule into molecule into metabolism . Together, these . Together, thermodynamics substrate pathways . requiresWork of the input work molecule, again molecule, according to the prin of precursors differentthe all into building product . There are forms many of energy: radiant, electri synthesis coffee processes, All cools down). biotic or abiotic, are the resultof conversionsor transfers of al and they energy, proceedways according oflaws to the Thefirst oflaw thermodynamicsenergythat states cannot be created transferred only or destroyed, or converted from ofso the total amount one form energy to another, the in universesame. the The stays always of second law thermo blocks, and for the synthesis ofsynthesis the and for blocks, blocksthe all into building Thus, by more including itself. biomolecules, DNA complex structurethe encoding of encodes ultimately proteins, DNA the structure ofthe components of all and organisms. cells Chemical Engines living seen that have or arethings and highly complex We ganized. assemble Theycomponents, own their synthesize and performthemselves ac other biological all functions, cordinginstructionsto the Synthesiz DNA. their encoded in and cells and maintaining and assembling biomolecules, ing organisms, are forms of energy whichenergythe is cal, mechanical and chemical energy, stored molecules. atoms together in the bonds holding in Energy can be converted and from one formanother, into can be it transferred doing work heat flowby (e.g., or by whena pot of ofcup or your water up, heats stove the on ses the conversionof one specific one specific ofciple structuralsubstrateand Enzyme complementarity: interactan exclusive, with lock-and-key fit. Theof thousands different reactionsenzyme-catalyzed area cell in organized series ofinto reactions called interrelatedcell's the constitute pathways characteristic fundamental another of Central livingthings. of constituents main on the acting pathways cells—nucleic proteins, carbohydrates and lipids—areacids, essentially of another indication livingall the same in things, com their aremon origin. Enzymes far more specific and efficient than efficiency and The greatman-made catalysts. specificity, diversityof con to simultaneously cells enzymes allows structofthousands differentfrom molecules relatively few precursor for exist pathways Enzyme-catalyzed molecules. the . ------catalysts , folded into a greata into , folded and the positions of and the positions , function as , function complementaryof fit globular enzymes proteinshelp to shape, support and Fibrous differenteachin enzymes Eachcell. type of enzyme cataly Catalysts areCatalysts that speed substances up chemicalreactions chemicalthe All process. the in consumed being without re ofactions a different enzymes, are a cell by one for catalyzed eachreaction, whichrequires presence the ofof thousands foreignor (antibodies), serve substances as messengers (hormones suchand growth as insulin hormone), among The other functions. and diverse most abundant many group of proteins, the globular and bind to many differentand bind to many dif and mediate many molecules proferent example, Motor proteins, for biological functions. structuresmany in duce movement includingcilia, flagella and proteins globular Other muscles. transport (e.g., substances defend againsthemoglobin,blood), in the oxygen-carrier of skin, scales, hair, nails and feathers, nails of proteinsthe collagen hair, scales, skin, of cartilage, bones, and cytoskeletonproteins, and tendons, which give structure and organizationto the cytoplasm of Most proteinsare cells. many variety of conformations, which enable them to recognize cules. Throughcules. these interactions, proteinsare able to carry instructionsthe out arebecause they key DNA in contained structures, cell all in components participants and key all in cell functions. protectand organisms. cells They include the keratinproteins Proteins initiate the assembly ofProteinsthe assembly initiate structures cell recogniz by to molecules, other with associating and spontaneously ing organelles, cells form supramolecular Ultimately, complexes. accord and organismssame way, the essentially in assemble specific interactionsto the ing of proteins mole other with translationof proteins,into and the specific interac mRNA oftions proteins whichmolecules, and other ac ultimately ofaspects all for count structurelivingin function and things. tions encoded in DNA areencoded DNA in tions carried out. The interactionsof accordingbiomolecules, to the their 3-dimensional structures groups, functional oftheir principle a fundamental is life. It appliestranscription to the of the molecules, RNA into DNA ticular amino acid sequence. Theacid sequence. amino ticular conformationspecific of a protein,of the specific positioning and groups,functional its to recognizeit allows and interact in other molecules with verya 3-dimen together in fitting pieces like ways, specific Throughpuzzle. jigsaw sional interactions,these instruc the 42 BIOL F150: Introduction to marine biology where the oxygen efficient zero. isnear where More concentration in the ocean fishes called coelacanths, deep primitive living forexample, fermentation, from isms still obtain theirenergy molecular oxygen. lacked atmosphere Some organ Earth's life,atatimewhen of used by probably was ancient forms respiration oxygen. type of less efficient This of absence fermentation the ATP forwork. biological respiration pathways converge into a common pathway,Catabolic called in all cells and organisms. work, biological all other typesof energy, usedin synthesis and ATP of is the common carrier todowork. energy the released phosphate and capturing down into ADP and by it back reaction, breaking endergonic phosphate) and phosphate. ATP an can then be used to drive goes into synthesizing ATP,reaction ADP (adenosine di from during an exergonic released the energy phate). Someof intermediate—usually a shared through reactions, to endergonic reactions exergonic from is transferred (C&HFig. cells,process In living 4.1). energy ( is an energy-yielding catabolism while process, ( components. is an energy-requiring Anabolism metabolism complex ones,simpler molecules and into more work. used toperform andcanbe molecules into simpler onesduring metabolism, complex by of is released the breakdown energy chemical molecules.This in organic stored energy into the chemical is converted sunlight, which of energy is the radiant energy of ultimate source engines. chemical Their things are Living the environment. light or heatfrom it absorbs when dered disor and usuallymore becomes maintain its organization matter, to energy on theotherhand,cannot useexternal within themselves.ating andmaintaining order Inanimate cre they are heat),while into simpler onesandby off giving (by downcomplex molecules breaking theirsurroundings of alsoobey thesecondlaw,They by theentropy increasing law. with the first in accordance work—all it into biological andchannel it into a useable form, the environment, convert from energy toextract things have the unique ability Living ( disorder or in the randomness with an increase proceed dynamics states that all processes , which breaks down glucose to supply most of downglucosetosupply mostof breaks which , breaks downcomplex molecules into simpler breaks ), breaks down glucoseincompletely,), breaks in the Anabolic metabolism Anabolic Anaerobic respiration Anaerobic entropy ATP (adenosine triphos ) of the universe. theuniverse. ) of endergonic synthesizes exergonic catabolic (or ) ) - - - - - many bacteria, it provides the carbon skeleton for every oth theskeleton carbon manyforit every bacteria, provides othermolecules. In of into a huge variety can be converted agoodfuel andit andtherefore things. in energy It is rich all living virtually of tothe metabolism Glucose is central issummarizedin thefollowingdives. equation: Respiration activity, long performing and seals when of asdowhales duringbursts respiration useanaerobic crocodiles sluggish Forficient oxygen example, normally respiration. for aerobic isinsuf there when periods forshort it to supply energy andcanuse respiration, foranaerobic the capacity retain however, Mosthigher organisms, cells and organisms. still ken down into carbon dioxide and water, is used by most respiration aerobic riods when ATPriods when by is not being generated the light phase pe glucose, during dark stored from energy to derive also userespiration Photosynthetic organisms respiration. through and tofueltheir processes, rials for their growth orindirectly, directly tosynthesizers, mate raw toprovide pho matter from onorganic depend All other organisms photosynthesis can be summarized in the following equation of process overall compounds.The cose andotherorganic phase ATP. of energy the chemical In the second phase,the into is converted which light energy,ecules absorb someof photosynthesis, called the of dioxide carbon glucose from andwater. phase In the first synthesizing of it into thework sunlight and transfer of ergy tosynthetic Only ingredients. inorganic molecules like glucose from synthesizing organic of capable are only someorganisms materialforbuilding other molecules,as afuelandraw use glucose use. Although almost all organisms for future starch molecules, carbohydrate like larger of housed in the form molecule needed by the cell. It also can be ware er organic C 6CO , ATP's energy is transferred into the synthesis of glu into the synthesis of ATP's, istransferred energy (in plants) and 6 H 12 2 +6H organisms are able to harness the radiant en theradiant toharness able are organisms O 6 (glucose) +6O light energy 2 O , in which glucoseiscompletely bro in which ,   glycogen C chemical energy 6 H light phase light 12 2 O

 (in ), and saved 6 i (glucose) + 6O

6CO 2 , +6H pigment

2 O O 2 pho mol dark dark ------43 Lesson 4: Biological Principles, Cells, Organisms & Ecosystems ------dy ho receive and . When changes. When environ their occur in fromfun another is environment the . Although they do not visibly change from visibly do not they . Although , whichof attribute fundamental another is life. open systems open ings, and respond appropriately. Theability to and respondings, appropriately. respond to signals damental property of Organisms living things. sample may surroundingstheir the availabilityfor of or oxygen light, presenceglucose, or the ofchemicals, or other or toxic pose a threat, or be ganisms food, may that compete for organa multicellular within cells Similarly, mates. potential information.exchange must ism and organismsCells must Homeostasis is achieved regulating by metabolism. of All the interconnectedcell's metabolicare pathways precisely regu respon coordinated and exquisitely a superbly lated into sive system. Eachof ofthousands the differentmetabolic productits supplies pathways at a rate matches exactly that for product,adjusting that demand for cell's the continually the changing needs of changing the cell, and for external Throughcircumstances. regulation,metabolism achieves is main and homeostasis efficiencymaximum and economy, tained. Livingarethings self-regulatingat every level, and respectwith every to aspect of Theinstruc functioning. their regulationfor tions are and are included DNA in carried out regulatory by primarily largeA proteins. portionof is DNA devotedproteins,these encoding much to and of workthe ofself-regulation into life goes homeostasis. and maintaining orderIn to regulateand organisms cells themselves, must information vital obtain continually about surround their logicalwork. absorb They raw continually synthe materials, breakand size release and molecules, down Because wastes. are exchangingthey energy constantly and materials with surroundings,their livingand organismscells are described as individual moment to moment, their areatoms and molecules Thebeing exchanged. constantly types of they molecules contain, and the concentration of remain each type, however, relativelyTheconstant. concentration of for cells, in glucose remain will example, nearly rate because the constant of its rate the balanced by just is ofor synthesis intake break its down or conversion product.another into Thiscalled a is namic steady-state and organisms is disturbed, cells steady-state ment and this order in make adjustments to return to a new steady-state. Thiscalled is steady-state ability a dynamic to maintain meostasis ------. We will learn will . We more about photo chemosynthesis and absorb energy, convertingabsorband and transferringit energy, bio into it In its composition, complexity and organization,and complexity composition, its In in exists life sharp contrastsurroundings. to its of Instead isola in existing livingarethings dependent absolutely on their however, tion, Livingenvironment. and organisms cells releasecontinually cess of and chemosyntheticsynthetic organismsmarine 6. Lesson in Self-Regulating cellular prokaryotescellular and eukaryotes. Some marine microbes, such as some bacteria living at great depth around hydro thermalorganic synthesize vents, absencethe matter in of Instead oflight. use inorganic they compounds solar energy, energysuchas an sulfide as hydrogen sourcepro the for Theorganismsphotosynthetic we are familiar most arewith half at least However, flowering ofthe plants. land photo the on Earthsynthesis vastsea, and the the conducted in is major ofity organisms marine photosynthetic are microscopic, uni again in photosynthesis. Sunlight is the drivingis again Sunlight force behind photosynthesis. in cyclingcontinuous this of among and carbon dioxide oxygen organisms, and between organisms their environment. and explore will these relationships when discuss we further, We some ofbasic concepts the oflesson. this later in ecology, whilegeneratesand respiration carbon diox oxygen uses ide. Thus, photosynthesizers and other organismsare in terdependent.provides Photosynthesis as the fuel as well required aerobic for the oxygen respiration. Aerobic respira turn,tion, in which generates carbon dioxide can be used matter, glucose. In respiration, In glucose. brokenis glucose into down matter, and energyreleased.is These are and water, carbon dioxide also reciprocal processesthe sense that photosynthesis in as a by-product, and generates oxygen carbon uses dioxide As the equations for photosynthesis and respiration show, and respiration photosynthesis for equations As the show, these are 4.5). In processes opposite essentially (C&H Fig. and water arecarbon dioxide sunlight, abphotosynthesis, sorbed from and used to create the environment organic ofcase ofthe In photosynthesis. photosynthetic humans, organismsare source ultimate the of energy for only not our machines,coal, petrosince also for but our bodies, naturaland leum gas wereformed all fromaccumula the oftion dead organisms countless of millions over years. 44 BIOL F150: Introduction to marine biology replication duplicating its DNA called molecules,duces byprocess a first by DNA and implemented bystructed proteins. A cell repro in ultimately molecular in nature, are again, processes, cells.These pre-existing cells from tissues, by new creating damaged cells and repair worn-out and continually replace anddevelop, alsogrow themselves. Multicellular organisms like individuals offspring, new can produce All organisms andmaterials, to is their ability ergy things, living however, their en andamajorinvestmentof of characteristic mostremarkable maintain homeostasis.The environments, and and external their internal signals from and act on receive and movement. They thesis, transport like syn into work, energy and convert metabolism through them theenvironment, transform materials from raw extract continually machines.They chemical bling and self-regulating unique and incredibly complex, self-assem things are Living Self-Replicating tosignals. andrespond cells receive living by which chinery to the molecular ma , can be traced schooling ments of move ocean,orthe perfectly synchronized miles of of dreds hun for locating a mateacross exampleresponses, awhale most complicated The at the molecular level. mechanisms bymediated are sight, smell,tasteandtouch, including proteins. All sensation, pendent on the specific actions of andis de atthemolecular level processes stems from andactoninformation things to receive living of the ability lifewehavediscussed, theotherattributes of of As istrue activity.and thus their biological their 3-dimensional shape tain and DNA limits, undergo proteins When these conditions exceedcer their surroundings. of pH andsalinity on the temperature, depends in turn which ontheir3-dimensional structure, ity tofunction depends abil Their fragile. DNAmacromolecules, and proteins—are things—the informational living most critical components of The signals forcells and organisms. important particularly theenvironment are and salinity pH of temperature, The tissues complexin a organism. different location among orby fuel al athreat, coordinating or food,or away from appropriately, respond light for example moving toward . It then divides into two . It then divides grow daughter cells denaturation and reproduce , losing , each , each ------. prokaryotic cells, and numerous chromosomes in the cells chromosomes cells,and numerous prokaryotic for the single DNA molecule contained in most chromosome complex called a into a supramolecular DNA is organized its cells. molecule of Each of cated in each DNA molecules is called its set of unique life.An organism's andperpetuates characteristics, and species these individual tion maintains the continuity of to the next genera Transmitting this generation. information the previous its DNA, inherited from sequencesof nucleotide in encoded the and asaspeciesare as anindividual ganism or characteristics, The (chemistry, physics, genetics), and by technologies. new scientific fields possiblemade bydifferent contributions from duringthetwentieth century, in biology advances important amongthemost were DNA andhowit replicates of structure the elucidation of DNA. The theparental containing a copy of formational sequences is truly incredible. DNA is incredible. auto sequences is truly formational these immense in things to preserve living of ability The all thetime. being added with more ous otherorganisms, fornumer DNA sequences havebeendetermined tire in biology, medicine and biotechnology.researchers En beingusedby tothepublic and already is available which genome sequencein 2003, human whole pleted thefirst com in 1990, begun genes. HumanGenomeProject, The 25,000 3billion comprising about nucleotides, about of somes in the human genome contain a combined sequence thousand genes.chromo 46 The several long andinclude million nucleotides rial DNA sequence may beseveral molecules. byDNAthe largest A bacte molecules are far stages. developmental at different occur in a cell or organism for that overgenethe time changes is responsible expression andfunctions.Changing structures them their different giving cells, types of in different off onorturned turned genes are different In multicellular organisms, in cells and organisms. all processes of is critical for the regulation and translation, transcription of the regulation through gene expression, Regulation of yield its product. translated—to proteins, of in the case DNA istranscribed—and of thatsegment when or RNA molecule. ther a protein for ei synthesizing one functional product, the information A eukaryotes. of gene is a segment of DNA which contains DNA which of isasegment genetic traits Gene expression genome , which identify an or , which chromosome , which is dupli , which occurs occurs , one ------45 Lesson 4: Biological Principles, Cells, Organisms & Ecosystems ------tro . Eco . Ecology . Thefull intraspecifi interspecifi (living)aspects of , and ecosystem , relationships. It begins physiological adapta biotic ecological niche (C&H Fig. 10.13). a com In (C&H Fig. . Theyinteract (C&H Figs. 10.14 10.15). and Figs. (C&H ecological succession of and physical populations, trophic population . Theofsystem interactions within habitat , with a large, with of volume autotrophs as food chain food food webfood (self-feeders).other organismsAll are (physical) and (physical) (feeding on others),on (feeding directlyor indirectly de community abiotic , with members, with of the same species livingparticthat in the interacting All species. other with an in populations autotrophs ofindividual organisms, growth,the and decline and trophic pyramid An autotroph is fed upon by a heterotroph, autotrophAn by fed upon is whichitself may another heterotrophic for become food organism, and so on, forminga in links ofmunity interacting many organisms, chains food are many a into interwoven relationships as 3-dimensionally be shown can also Feeding a base and increasinglythe fewer heterotrophshigher at is the study ofthe study is these interactions. all these interactions All result includingchanges, in the tion evolutionary adaptation a result, As changesenvironment. the in arecommunities The changing. replacementconstantly continual of one com called is another by munity interdepenaresystems many with and dynamic, complex dent partswhose interactions reshapecontinually the whole. level,fundamental most the At be viewedcan ecosystems termsin offlow the of from organism energy and matter, to organismand between organisms and the environment. Energy and materials flowfrom organism to organism ac cording or to their feeding, chemosyntheticand photosynthetic with organisms, which are heterotrophs pendent on autotrophsenergy their for and raw materials. Interacting of attributes unique the have outlined We living and things interactliving that seen continuously things envi their with and ronments, extractingand releasing energy and matter, responding of to a myriad Organisms signals. interact with the both or environment, their cally ular area,a constitute that cally areaform a and between the organismsand their a community, biotic an constitute environment, physical range ofecological characteristics of organism an or spe cies (includingspecific habitat,its reproduc feeding habits, etc.) is called its tive strategy,

. ------evolution evolution natural selection is produced. is Through mutation , are passed from generationto gen of time. Theover species modern theory ofoccursthat material genetic during the fundamental characteristics fundamental the forms all shared by of life. evolutionof firstits life since appearancethe geological in recordyears4 almost billion through changesDNA in ago, whichwere transmittedgenerations,to subsequent accounts both the immense diversityfor and of livingtoday, things win's hypothesis, demonstratingthat hypothesis, win's advantageousfor traits genetic results the in ary adaptation ofevolution conceptunifying become the has of biology, much theoryunifying the is as plate tectonics The of geology. heritance washeritance Charles known, first Darwin "the hypothesized survivalof selectiveunder fittest the pressure"ex as an changing the for planation appearance of time over species oforigin and the overwhelman then, new Since species. ofbody ing evidencesupport in accumulated has of Dar ent would confer advantagewould an ent organism. on that would It be more to survive likely and reproduce,al its passing tered advantage and the DNA, confers, it offspring. onto its before and the natureIn the mid-1800s, DNA of in genetic however, changes in DNA are changesDNA in beneficial. Thisoften occurs however, under challenging or changing conditions. environmental if example, For a previously availablemol fuel or nutrient ecule became unavailable a mutation the environment, in whichorganismenabled an a differentto use or nutri fuel due to the accumulation of accumulation due to the and to the DNA, in mutations recombination reproductionsexual eration.areMost mutations harmful, resultingdefectivein organismsto reproduce unable Occasionally, themselves. Evolving Species are altered vast over spans of time, as changes almost 4 billion years!almost 4 billion an unrepaired When error does in a portionof frequently occur, nucleotide the is sequence changed, and a genetic such the structure changes, of fi the for also allows DNA characteristicdistinguishing nal of life—its matically repairedmatically to and continues specific enzymes, by generation in replicateafter generation of itself, cells, Some formsof error. without almost been bacteria have reproducinggenerations countless for themselves with remainingunchanged for essentially nearly perfect fidelity, 46 BIOL F150: Introduction to marine biology the food chain, and eventually returned tothe environment and eventually returned the food chain, matter, into organic passed along ganisms, incorporated by dissolved in the ocean is absorbed photosynthetic or bon cycle (nutrient) cycles in for life - nutrients - occurs necessary matter types of different of recyling The forms. and inorganic and theenvironment, in organic ism, andbetweenorganisms used overcan be and over, to organ organism from passed forto continue. life Matter,is necessary on theotherhand, solarenergy, new therefore, continual input of The ronment. it is dissipated into the envi that originally entered energy all the in the chain, all the organisms and decomposition of andusedby things. living Eventually,captured with the death re be can never up thefood and passed chain, converted timeitis thatislosttotheenvironment, each energy The pyramid. the trophic of higher level at each consumers of biomass adecreasing tosupport required are producers of Vasta hugeor thechain. numbers, at theendof totheorganisms available would be energy only oneunit of by absorbed the producers, solarenergy units of 500,000 into the environment. So,links and with five in fooda chain is dissipated as heat link in the chain, successive at each energy, available of other90% The so onupthe foodchain. and consumers, tosecondary that is available of 10% about and only consumers, to primary is available thetic organisms by absorbed photosyn thelight energy of Only 2% about to the next. the food chain one link of from and transferred energy into chemical is converted producers, system through the enters Solar energy and irreversible. is unidirectional system, or theliving through energy of flow The matter,organic levels.on dead at all trophic organisms from Decomposers are feeding habits and carnivorous herbivorous top carnivores vores are consumers andhigher trophic-level Secondary and soonupthe food chain. consumers, on primary feed ers are onproducers directly that feed ers producers phic (or

levels ( predators herbivores , in which carbon dioxide carbon or in the atmosphere in which , , and heterotrophs are its are , andheterotrophs (C&H Fig. 10.16). Autotrophs are acommunity's are (C&HFig.Autotrophs 10.16). (or ( top predators ) and those at the top of the chain are are the chain ) and those at thetopof saprotrophs . We are already familiar with the We. familiar already are or grazers ). ) are consumers that feed consumers ) are Secondary consumers Secondary ). Organisms with both ). Organisms consumers primary consum primary biogeochemical omnivores biosphere biomass . Consum carni car ------, . ,

environment, much of the dead organic matter in the ocean organic thedead of environment, much to the terrestrial carbonate shells and skeletons. In contrast carboninto calcium ahuge amount of incorporate organisms matter, In addition to fixing carbon into organic marine Earth. fixation thecarbon (and oxygen on production) half about ter. for responsible are Marine photosynthetic organisms mat organic tomakemore bycan beusedagain producers matter,organic downdead break nutrients which releasing all continuation of alsois critical forthe decomposers of role materials.The andraw forthat can beusedby energy all other organisms carbon (CO inorganic of the conversion for responsible critical because they are is cycle in the carbon photosynthetic organisms of role The (C&H Fig.be used by 10.20). photosynthetic organisms it canagain where dioxideas carbon respiration), (through growth of individuals and populations, and affecting whole whole and populations, individuals and affecting of growth nutrient may a become that of the essential nutrients, then the availability any of of at any point in the cycling occurs, cycles.When abottleneck in similar biogeochemical and silica, participate as iron such by organisms, othernutrients required (C&H Fig.10.22).The in the and passedupthe foodchain matter into organic incorporated by absorbed producers, canbe in theoceanasphosphate,which available erally acids, is gen nucleic of ingredient anothercrucial phorus, in the vital role this perform bacteria and other microorganisms) types of used by producers. tobe in order that must be converted in forms ronment in the enviteins, is another nutrient commonly occurring acidsandpro nucleic an essential component of Nitrogen, caused by carbon dioxide increasing emissions. warming and global effect the greenhouse for moderating important pump to asthe shells and skeletons,is sometimesreferred into calcium carbonate andincorporated photosynthesizers byfixed carbon, marine amounts of vast movement of ward ervoir it is on the bottomwhere ending up in sediments decomposers, of reach sinks out of or of unavailable carbon and other nutrients. This down and othernutrients. carbon This unavailable of carbonsink nitrogen cycle nitrogen nutrient cycles Nitrogen-fixing . These processes havebeencritically processes . These limiting resource limiting sequestered (C&HFig.Phos 10.21). , andall life, since they organisms (certain (certain organisms 2 phosphorus cycle phosphorus carbon fixation , an enormous , anenormous ) into organic forms forms ) into organic , restricting the , restricting carbon res — ------

47 Lesson 4: Biological Principles, Cells, Organisms & Ecosystems

. , ------or in nu flow ben and litto zones. zones. ( obligate parasitism or , abyssal bathypelagic sexual reprosexual aphotic osmoconform , , , intertidal or . A food webfood shows . A and zones (Fig. 9). We 9). We zones (Fig. (bottom-dwellers)live (active swimmers)and facultative bathyal zones and, according to . In the next lesson, we will lesson, next the . In ), commensalism : mesopelagic nekton . Thespecies inter intimate most Benthos dysphotic oceanic , , hadopelagic osmoregulators and sublittoral . In Lesson 2, Lesson we organized. In ben the asexual reproductionasexual cycling of matter photic symbiosis and . Mutualism may be may . Mutualism ( , dependingtemperatureon regulation. diagrams,carbon cycle the for those like epipelagic hermaphrodism (animals living (animals on top of the sea floor) or neritic and pelagic (driftersand weak swimmers) live epipethe in ectotherms, endotherms, poikilotherms interspecifically , or subtidal (those within the sea floor),the within (those among others, and di zones (Fig. 2). In Lesson 3, we divided the pelagic zones (Fig. and mutualism depending on how they control dependingthey on how internalconcentrasalt ), epifauna homeotherms according distinguished be Theyto reproductive may strat example, for egy, duction learnmore about some ofrelationships, these as we move to Parton ThePlayers: Survey A Two, of Marine Organisms. habitat into depth, into abyssopelagic also organizedpelagic the according environment to il into lumination, we can organize marine organismsaccording Similarly, to habitatand lifestyle. while habitats, benthic in plankton lagic. course,this Later in further we will classifybenthos as fauna 334). 15.2 p. categoriesFig. (C&H size into vide plankton ex can categorize organisms For other ways. many in We be may ample, they ers Or tions. species, species, in occur actions and interactionsAll be reducedcan ecosystems in to the of energy the flowof The energy and matter through a community. "big picture" of cycledcontinually matter is between how organismsin be shown can environment and the trient cycle 10.20) and nitrogen10.21). cycle(C&H Fig. (C&H Fig. The two divisions, contains marine environment thic habitatthic according to depth, into ral hadal

. , ------su and fam ) and ) carni genus . Simi tissues , second , food webfood (Fig. 10). (Fig. organ sys , or with other , or with order, class, order, decomposers producers (in eukaryotes) (in whichto bind ( (C&H Table 4.1). (C&H Table omnivores organism (C&H Table 4.3). (C&H Table , and , and all the popula the , and all ), atoms for us), according us), for to the

organelles domain ). We can further can ). We order them , autotrophs ecosystem primary consumers grazers intraspecifically , building-block molecules building-block ( binomial nomenclature population , sapiens

and , whichcooperate in top carnivores (C&H Figs. 10.13, 10.14 10.15).and Figs. (C&H relationships chain, a food in ), . Macromolecules into assemble consumers , etc. (C&H Fig. 10.15). Or we can call them 10.15).call can Or we Fig. etc. (C&H , complexes Homo (

organs ( these . Communities interacting. Communities physi their with herbivores molecules . Cells (in complex eukaryotes) complex (in . Cells form predators ( , which make up the individual cells species then into progressively into then larger groups: primary producers trophic pyramid pete with their own kind, own their pete with We can show can We or Organisms Theyinteract com ways. many in communities in heterotrophs into ary consumers producers, vores community form cal environment an Organismsparticipate at differentcommunities in trophic Therelevels. (feeding) are phylum, kingdom Organismsof species livingsame the a particuin lar areaa constitute livingtions and interactingan areatogether in form a Eachof type organism its by can be identified and worldwideof system lar genera (plural ofare genus) groupeda into ily, pramolecular and which build tems Organismsare from assembled into gether macromolecules We have seen that organisms have seen that whileand ecosystems, in We arecrediblycomplex, governedareand principles basic by organized.highly review Let's some of we have ways the learned (here 4 and 10) representto C&H Chapts. and in organizationthis make sense ofand complexity. this all communities. Other essentials, such Other essentials, warmth as light, communities. or resources become limiting space, also when shortin supply. Schemes Organizational 48 BIOL F150: Introduction to marine biology sary at the back of your textbook. of attheback sary theglos guideandinyour textbook, andcheck thematerialinthiscourse re-read theseterms, any of themeaningof about unclear you're If for understanding. for thislesson.Doaself-check keyListed belowandonthefollowingterms page are • • • • • • • • • • • • • • • • • • • • • • adaptation adaptation andphysiological evolutionary mutation, selection recombination andnatural and genetic trait gene, genome, chromosome, gene expression andasexual reproduction sexual reproduction photosynthesis andchemosynthesis (fermentation) respiration andanaerobic respiration aerobic homeostasis ATP endergonic andexergonic lism andcatabolicmetabo metabolism anabolic catalyst, enzymeandsubstrate anddenaturation conformation DNA andreplication of translation transcription, systemandorganism organ tissue, organ, unicellular andmulticellular andeukaryote prokaryote elle andcell complex, organ membrane, supramolecular (DNA, RNA) andnucleicacid carbohydrate, lipid,protein, glucose, acid, amino acid, andnucleotide fatty building-block moleculeandmacromolecule compound compoundandinorganic organic andbiotic abiotic glossary terms - - • • • • • • • • • • • • • • • • • • • • • • • facultative andobligate symbiosis facultative symbiont andhost parasitism symbiosis: commensalism,mutualism and competition andinterspecific intraspecific succession ecological population, community andecosystem niche habitat andecological ily, genus andspecies domain, kingdom,phylum, class, order, fam binomial nomenclature andhomeotherm poikilotherm andendotherm ectotherm andosmoconformer osmoregulator benthos, planktonandnekton limiting resource andglobalwarming effect greenhouse carbon sequestration/pump/sink carbon fixation fixation andnitrogen cycle andnitrogen carbon cyle biogeochemical (nutrient)cycles biomass of biomass andpyramid pyramid andtrophic food chain,web troph) and decomposer(sapro omnivore carnivore, toppredator/ consumer (carnivore/predator), secondary consumer(herbivore), primary sumer) (con andheterotroph (producer) autotroph - - - - 49 Lesson 4: Biological Principles, Cells, Organisms & Ecosystems in your in your . For example, if example, . For asked to complete , but also , but clear and brief

• (Please make your answers and concise) complete your make (Please and submit the written assignment to UAF eLearning to UAF assignment written the and submit according Education & Distance to directions found Written Assignment LessonWritten 4 for habitat and ecological niche autotroph and heterotroph benthos and nekton and genome gene anabolic and catabolic pathways photosynthesis and respiration prokaryote and eukaryote osmoconformer and osmoregulator osmosis and active transport ectotherm and endotherm intraspecific competition and interspecific competition population and communitiy amino acids and nucleotides conformation and denaturation , ofnot those or Course textbook the Guide. Be

cover sheet m. n. i. j. k. l. e. f. g. h. a. b. c. d. that each of in the biosphere. these groups plays Describe three or strategies modes organisms used by to accomplish reproduction. Symbiotic relationships are the most intimate type of relationship Describe the three between species. types of give of own experience, an example symbiosis and, from your each type. photosynthesizers,Without nitrogen-fixers and decomposers, life on Earth would grind to a halt. Explain the vital role The of attribute is a fundamental ability to self-assemble and briefly do this, how they Briefly explain living things. describe three of other unique attributes living things. Define the following pairsDefine the following of also include but information. all the essential terms. Comparing/contrasting Be brief, the example: two terms For to do this. will often help you atoms, carbon more 2 or of a "backbone" of consist compounds - organic compounds inorganic and organic "backbone." a carbon lack compounds Inoganic oxygen. and hydrogen as such atoms other to bonded 3. 4. 5. 2. 1. "describe" something, do so - don't just list. (To be clear and show your understanding, be clearyour and show are imagineto you that explaining (To list. just do so - don't "describe" something, someone who of has no knowledge the subject.) Includea and in the course received.online at eLearning.uaf.edu materials you Study the information presented for this lesson until you feel that you have a good understanding. Write your answers your information the understanding. have a good Study Write feel that you presented you until lesson this for own words 50 BIOL F150: Introduction to marine biology Part two The Players: A Survey of Marine Organisms 52 BIOL F150: Introduction to marine biology 53 Lesson 5: Microorganisms And The Tree of Life - . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). ("Evolutionary Perspective"), pp. 79-83 ("Classifying Living Things"), p. 216 ("Eye on 216 ("Eye ("Evolutionary 79-83Living ("Classifying Things"), Perspective"),p. pp. Science: The Census of 218-219 ("Symbiosis") Marine Life") and pp. Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Recognize the greatRecognize abundance, ubiquity and diversity of microorganisms. Understand the importance of microbes to every aspect of life on Earth, and recog nize our lack of of knowledge them. Become acquainted with some of the marine microbes, representing the three major divisions of life. ofBe familiar with the Tree Life, and how it is a work in progress. the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education. C&H: Chapter 5 (pp. 85–100) and pp. 349-350 ("The Microbial 73 and review Loop") 85–100) and pp. C&H: Chapter p. 5 (pp. More information, including interactive flashcards, quizzes, videoclips can be and links, 3. 4. 5. 6. 7. 1. 2. Reading assignment Reading steps for completing the lesson the completing steps for Objectives of this Lesson of Objectives And The Tree of Life And of Tree The In this lesson, we begin a survey of the forms of life, starting marine with the smallest, but least significant: microorganisms. no means by Microorganisms Microorganisms 5 54 BIOL F150: Introduction to marine biology I tional way to study microorganisms has been to tional way tostudy microorganisms tradi The afossil record. of and they don't leave much these useful organisms, for classifying too simple to be very are made visible under when the microscope, even tures, struc Microbial and animals, andtheir detailed fossil record. complex beendoneby things,living has traditionally looking at the isms so difficult. make studyingmicroorgan and classifying factors Several will continue to evolve. life in the future of the tree we draw How technologies. new of often with able, the emergence avail becomes information asnew cess—subject to change science as anongoing pro texts is a primeexample of biology life in recent of the tree recently. of redrawing The lopsided,until quite other) has been very toeach related Life ible the universal andaccessible life.As a result, vis more behind that of far has lagged microorganisms of studying them, our knowledge of the difficulty Because of The Tree of Life on Earth.. organisms in and, many ways,the mostimportant ever, diverse the mostabundant, are that microorganisms isshowing us,how Recent them. we knowabout research howlittle of them in one lesson,issimplydescribe measure a into one category, organisms andattempting to these diverse life, of division thethird of est members include Archaea of allmembers include They not all related. are cant: with the smallest,but bymeans leastsignifi no starting marinelife, of the forms of a survey In this lesson, we begin (the branching diagram depicting how living things are things are how living depicting diagram (the branching single-celled algae morphology , two of the three major divisions of life. They also life.They of majordivisions the three , two of microorganisms Microorganisms and the tree ofMicroorganisms and life Taxonomy (structural characteristics) of plants of characteristics) (structural . Microorganisms, or . Microorganisms, and , the practice of classifying classifying of the practice , protozoans Eukarya (A Huge Cast of Tiny Characters) (A Huge Castof Bacteria . Combining . , the small microbes cultivate Tree of Tree of and • ------,

vive and proliferate—about as likely as winning the lottery! aslikely aswinning the lottery! andproliferate—about vive light, acidity, pressure, to sur unique organism foretc., each temperature, and at the right conditions of concentrations, medium, in the right nutrients in the growth combination of the right onproviding depends process this laborious of success The culture." those typesin a "pure of each tivate thesample,andthen cul out of that grow microbe types of isolate allthedifferent nature, samplestakenfrom (grow) be determined in a much more accurate and meaningful accurate more in a much be determined can so their relationships organisms, of genetic information the analyzealso allows scientists to directly and compare without culturing isolating or them. Molecularsequencing microbes, gies made it possible to detectandcharacterize technolo in length). These nucleotides millions to billions of (remember,databases DNA sequence is an theorganism's hugeto analyze needed such were which available, came be DNA molecules. At the same time,powerfulcomputers in RNA nucleotides and of molecules, andthe order protein amino acids in of the order allows scientists to determine late 1960s, all this. havechanged In the technologies new Revolutionary plex otherandwith other organisms. associations,with each since they usually nature, exist in intimate and com their true studying us from may isolating microbes prevent of practice At the sametime, the is moisture. there especially where thing on the planet, and nonliving living in and on every living everywhere, are since microbes problem, is another samples many marinemicrobes.Contamination of to capture coarse too are century than a that scientists used formore ocean. and unevenly distributed in the vast sparsely can be since microbes lecting problem, samples isa col of In marineenvironments, theprocess even cultivated. have been found in nature microbes of Only a tiny fraction molecular sequencing was developed, which which developed, was Plankton nets ------

55 Lesson 5: Microorganisms And The Tree of Life . . ------Glob Archaea Eukarya , and (Fig. 11 and C&H (Fig. Archaea

, commissioned by the by commissioned , ,

animals plants protists fungi bacteria Bacteria of life: 270,000 80,000 72,000 4,000 Domains 3-domain phylogenetic system 3-domain phylogenetic Fig. 4.24) has become widely adopted. Thebecome widely 4.24) has ofnumber Fig. king each doms within being determined. domain is still At least three major groups of archaea, and several dozen major groups ofDoman Eu bacteria, have been identified. Within karya,it remainand fungi plants but animals, as kingdoms, unclear still is to classify how diversethe eukary unicellular tree A protists. the otes, ofthree life with branches, main two of which prokaryotes, are occupied by representsa at the highest taxonomic level,taxonomic highest at the organisms and do the within eachgroup share a common ancestry?most Have we found of forms the of life on Earth?recent in Discoveries decades have provided some surprisinganswers questions. to these and coworkers Carl1970s, the In Woese se used molecular to comparequencing nucleotide RNA bacteria, sequences in Severalwerespecies as discovery. and made astonishing an different from were the other bacteria as they from eukary Theseotes. organisms, unique producers methane in found swamp were muckintestines, and animal named moreAs oforganismsthese were discovered analyzed, and became clearit previouslythis that unrecognizedgroup requiresit that so unique is own branch its on the tree of proposedrank taxonomic an additional 1990, life. In Woese abovethe three to distinguish Kingdom, dif fundamentally ferent This With 4 ofout With 5 branchesdevoted to eukaryotes, tree this are eukaryotes. far, by living most that suggests things, proportionsSimilar are evidentwhen we look at the total numbersof The identified. have been that species al Biodiversity Assessment Program Environment Nations United 1995, in reported: 1,320,000 More the yearsidentified in species, be added to since, may the proportionsthese totals but remain roughly the same. Do proportionsthese reflect proportionsactual the of livingthings Do these five groupsexistence? in deserve all to be ranked . ------name, , orders Animalia orders species , was proposed and for the mushrooms, for cell seen in bacteria, in seen cell and , but has also radically has , but washy His published. cell found in plants, ani plants, in found cell Protista Plantae (or divisions), and phyla divisions), (or phyla and Fungi genus 5-kingdom phylogenetic 5-kingdom phylogenetic , familiesinto eventually became unify the eventually (evolutionary history, or relat (evolutionary history, phyla prokaryote eukaryote families microbiology phylogeny . When Carl. When (a.k.a. Linné Carolus Linneaus) , classes into for the bacteria, and for The Origin of Species natural selection , in use since the 1950s and still found in many many in found 1950s the since and still use , in (heterotrophs that absorb organic material) and heterotrophs structures and lifestyles Monera/Bacteria—unicellularprokaryote autotrophs Protists—mostly unicellular eukaryotes,Protists—mostly unicellular diverse with eukaryote multicellular Fungi—mostly osmotrophs Animals—multicellular eukaryote heterotrophs Plants—multicellular eukaryote autotrophs classes kingdoms Monera

textbooks, produced a treetextbooks, offive life with branches: main molds and yeasts. Theweretypes also cell basic two and yeasts. molds recog simpler smaller, nized: the morethe and complex This and protists. fungi, mals, system ing things. Phylogenetics was whatbased on still Phylogenetics be could things. ing observed time: morphologyat the record. and the fossil centuryAnother passed before two morewere kingdoms split out: pothesis that species evolvespecies that pothesis through time over prothe cess of concepting ofdirected It to become a taxonomy biology. search the for edness) of their place the naturalin species, order of liv A century later, a third kingdom, centuryA later, the diversefor group oforganisms unicellular did not that and Charles seem or category, to fit either the plant animal Darwin's then groupthen them into into into only introducedmid-1770’s, the in system taxonomic this weretwo kingdoms recognized: way. Applying techniquesthese microorganismsto not has way. revolutionizedonly altered our viewof relationships the livingamong all things. organisms identify their by We 56 BIOL F150: Introduction to marine biology tinction causing a are thathuman activities It appears is decreasing. andglobalbiodiversity rate, at an alarming disappearing At the same time,weknow that species are marinespecies. of and91% the totalspeciesonEarth, of then we haveyet todiscover 86% this is accurate, marine. If are which million to be8.7 species,million 2.2 of biodiversity studymillion to 100 species. estimates global One recent million 3 wildly from ranges estimated by taxonomic experts than they Global biodiversity can be describedorcatalogued. faster all the time, much being discovered species are New is: We them,the answer don't know.er wehavefound most of and wheth are, how many species there As to the question of than do eukaryotes. rapidly more forms they new evolve freely, andexchange genetic material more rate, faster so at amuch also replicate existence. Prokaryotes Earth’s of all that time, formost and diversifiying have beenevolving than multicellular plants and animals!They earlier years billion 3 andabout than the unicellular eukaryotes earlier ago, 2 billion years about billion 3.8 years all, they arose life.After of should up asignificant make portion karyotes things.among living It makes sense,however, that pro and relationships life on Earth of view different radically of proportions unprecedented in the history of life of in the history unprecedented proportions of Figure 11. A simplified version ofthe Tree ofLife, following the3-domain system. mass exmass - - - per 200 m of the ocean holds an estimated5x10 theocean m of 200 per in the ocean is astounding. up microbes The abundance of at altitudes as high miles.as 50 floating The found airborne, be caneven ontheplanet.They substrate every saturate bodies, and fill the oceansandfreshwater They abundance. in great everywhere, discovering,are we are Microbes, Microbial Abundance Abundant, Ubiquitous Diverse and this out. cover them.Whatlittle we doknow bears maymany wecandis unique microbes belost before but mostly unknown to us, and that lific and diverse, lifein the sea is pro all suggests that microbial This they discovered? canbe being lostbefore are many more disappearing,then how known species are (C&H p.If 215). karyote abundance in both saltwater and freshwater bodies. and freshwater abundance in both saltwater karyote thetotalpro of environment, 99.8% cells in the pelagic this adds up to 10 is takeninto account, theocean When theimmensevolumeof 5x10 toabout cells drops of m, the concentration 200 water. thimblefulBelow of cells in every prokaryote 500,000 29 (1 followed by 29 zeroes) prokaryote (1followed by prokaryote zeroes) 29

5 /ml, or /ml,

4 /ml. Illustration by Christen Bouffard, UAF eLearning UAF Bouffard, Christen by Illustration - - - - 57 Lesson 5: Microorganisms And The Tree of Life - - - - (“extreme-loving”). . Thisarchaean, col Strain 121 extremophiles —like it really hot, also found in hot really in it —like hot, also found —found in extremely cold environments, like extremely cold environments, in —found —like it hot, found in hot springs and aroundsprings hot in hot, found it —like —prefer extremely high pH conditions, found —prefer found pH conditions, extremely high —grow best under extremely low pH condi (or piezophiles)—live under extreme pressure, —live in extremely salty environments, deep-like —liveextremely environments, salty in —those livingdeep beneath the Earth’s surface, alkaliphiles and alkaline soils. soda lakes in carbonate springs, so far Most extremophilesfound are ar bacteria. Many chaea are also including extremophiles, record-holdingthe hyperthermophile, called lected from hydrothermal de Fuca Ridge, Juan at the vents nearly Ocean, survivedPacific the 1.5 down in miles and hyperthermophiles springs and hydrothermal and on active vents, seamounts, where lava is emitted directly onto the sea floor. psychrophiles cold the and sediments, sea ice Arcticand Antarctic seawater, deep sea, and lake ice and glaciers. barophiles such as deep-sea and subsurface environments. halophiles sea hypersaline hypersalinebasins, Dead Sea the like lakes and the Great and solar salt-evaporation Salt Lake, ponds. acidophiles at some hydrothermal found and springs hot tions, vents, mines. Extremophiles MicrobesEarth'sthe beneath miles areonly not found being everysurface,in but extremeimaginable, environment so suchwe call microbes Extreme from only of our point these environments view, are normal perfectly organisms the for living there. fact, In survivecouldn't most anywhere Extremophiles else. include: endoliths at extremetemperatures or oxygen, light without and pres sures, and inorganic feeding on hydrocarbons compounds, some reproducing once every only 100 years. thermophiles hydrothermal vents.

. ------cyano biomass epifluores picoplankton , was discovered . These . becoming clearIt is mi that subsurface prokaryote organ 30 Prochlorococcus biomass in the late 1970s. When treated When 1970s. late the in with Synechococcus called isms. Most occur beneath the oceans, constituting perhaps constituting oceans, the occur beneath Most isms. one-third of Earth's croorganismsand beneath the sea in are the most abundant organismsEarth,on arethey and us. to largelyunknown plains extend as deepextend as 4 plains km (2.5 mi). Therep volume this resentsso enormousis concentrationsthe that, although of microbesofnumber total the herebe less, may subsurface microbes is greatersediment or the water. in shallow in than There are an estimated 10 microorganisms years, many for been known has from for mation fluidsof have studies oil, gas Recent and water wells. prokaryotesthat shown are and often abundant, ubiquitous aquifers, petroleum deposits groundwater, in and sediments. underlyingSediments ofmost ocean floor the coastal the and Since the sea floorSince 70% offor accounts the Earth’s surface, representsthis large almost unimaginably numbers of mi croorganisms Themarine sediments. in greatest numbers be the waterofnot in may or on the microbes, however, deep The beneath it. but of existence ocean floor, subsurface otes account for morefor otes account 90% than of the ocean’s Microbes are even moreocean floorthe on abundant in than the waters above.con and soils marine sediments Shallow abouttain 1000 more times prokaryotewater. in than cells ters, a common occurrence now is Cells, “Tiny (C&H p.92, Surprises”).Big appearsIt prokaryote that members of far plankton the eukaryotes, outnumber microscopic the algae and protozoans previouslyto dominate the thought bacteria, archaea and microbial eukary Together, plankton. appearedorganismsabundantmost the to be Earth,on a related until genus, 1988,in was that even morenew Finding abundant. types ofarchaea,and bacteria great in pelagicin abundance wa In 1979,In a newoftype microorganismwas reported using of a genus epifluorescence miniature microscopy, bacteria categorysize (the livingsmallest the were plankton) for they ocean that the in numerousdistributed so widely and Many ofMany microbes marine these are es they that so small capeddevelopmentthe until detection of the microscopecence microbialfluorescentultravioletunder glow cells dyes, light instrument.this with counted and seen be easily can and 58 BIOL F150: Introduction to marine biology including nutrients, light (C&H p.including defenses, and even 90). services of with variety a ists may theirpartners provide for mutual their existence. Microbial the host or partner symbiosis. in engaging opportunistic, merely Some are win-win situation. a biological that benefit both partners, form the host. Still others of sites norhelp their that neither harm form symbionts kind of may in every found be engaging microorganisms ing world, the liv Throughout onEarth. substrate living also every but substrate, nonliving not only every saturate Microbes Intimate Relationships ft underground! anancientnow 2,000 sea bed tals from in amber,tombed in salt crys million 250 years and after en bee a glacier, in the gutof million years 25–40 after in a buriedft deep 300 years 750,000 after and revived andhavebeenfound for years, unprotected the Earth orbiting after in a spacecraft, havesurvived rial spores time.Bacte but also forseemingly indefinite of periods any conditions, not only in virtually survive Microbes light or oxygen. or toxic conditions, of andin the absence acid,alkaline orextremely but also in the presence sure, and pres temperature of not onlyunder extremes thrive vents hydrothermal at deep living Organisms Sea. ranean halophiles oceans, in the and cold deep living and archaea polyextremophiles vents. Manyare extremophiles hydrothermal around living alsoexist, Multicellular like the animals extremophiles inch. 8 tons per square of in the Mariana Trench, at pressures shelled some soft- extremophiles, including also are ote microbes Someeukary and spores. cient to kill all microorganisms thought suffi previously at121°C, autoclave pressurized minutes is 15 in a conventional sterilization regimen The (250°F). at 121°C cultured when doubledeven its numbers or commensal foraminifera symbiosis pathogens Obligate live in, the bodies of larger organisms. Some organisms. larger of in, thebodies live in the deep hypersaline basins of the Mediter basins of hypersaline in the deep symbiotic relationships, neutral guests neutral symbiotic relationships, . symbionts, on the other hand, require symbionts, on the other hand, require , benefiting themselves at the expense , such as , such Ectosymbionts pulled from the pulled from psychrobarophilic mutualistic hosts live on,and live . Others are are . Others Challenger Deep relationships relationships facultative psychro bacteria endo para ------

forlife in these lightless, worlds. sulfur- andmethane-rich harbor andmussels, clams tube worms, communities,including animal life in these rich vents and coldseeps.The thermal hydro communities around highlyonts drive productive symbi Other microbial foraminifera. microscopic and even anemones, andsea as well as with fans sponges,mollusks sea like soft corals, reef-corals, of sociations with relatives similar as Otherzooxanthellae form and ahomein return. nutrients inorganic nutrients, receiving andorganic energy called endosymbionts,commonly These reefs. coral of formation gellates involves photosynthetic partnership, One such and dispersal. forment, nutrients,reproduction route a or shelter and astable environ they may receive In return, while pathogenic microbes mediate disease,in all organ pathogenic microbes while essential to life, processes mediate tic microorganisms mutualis and varieties, life.In untold numbers of fabric essential curiosities,bonds, wovendeeply but into the are not mere are relationships symbiosis. These of importance and the pervasiveness We only to recognize beginning are all examples of are roundings. These prey,municate, lure matesorblend into their sur attract some fishes,octopuses andsquids, enabling them tocom or ria colonize specialized light organs, Luminous bacte source. any additional carbon ornitrogen wooddiet,without on a pure to survive allows the shipworm enzymes for cellulose This digestion and nitrogen-fixation. cellulase and nitroginase, endosymbionts provide These also house bacteria in specialized glands within their . wood-boringclam) (actually atypeof function. Shipworms this dosymbionts in embedded specialized cells to perform with en reduced, or greatly also is absent digestive tract the vents and coldseeps, hydrothermal around living valves and bi symbiotic In bacteria. tube worms chemosynthetic filled with entirely system,but instead are or excretory they have no mouth, gut, anus become gutless wonders: for example, have worms, oligochaete sediment-dwelling ways. Some in remarkable the participants transforming time, spansof overvast develop Symbiotic relationships zooxanthellae chemosynthetic of the genus of , provide reef-building corals with corals reef-building , provide Symbiodinium microbes which provide thefuel provide which microbes obligate , which drives the drives , which symbiosis. dinofla , in ------59 Lesson 5: Microorganisms And The Tree of Life . , ------. Biofilms biofilms .” Microbes are quorum sensing chemical chemical signals marine snow everyMarine microbeshull. ship’s . When the levelthe . When of autoinducers in foul (C&H Fig. 5.4). Fossil stromotolites can be 5.4). Fossil Fig. (C&H autoinducers called a particulararea bacteria beginsufficientlythe is high, to form a biofilm. Thisprocess called is Microbesproduce biofilms in inhibit that other molecules the quorumof sensing competitors. their Eukaryotes, for also produce chemicalsmay some seaweeds, that example interferequorum with thereforeand sensing prevent fouling ofservethemselves Biofilms biofilms. surfaces their by as are complex, dynamic communities ofarecommunities dynamic complex, interactingorganisms. They manufacture, transport, re and compete for consume, sources,via and communicate corrodeequipment medical pipes and contaminate metal air and heat. They and the systems that deliver our water, cling shower curtainto the bowl, cause and toilet and the giverocks.wet on slip to us us they teeth, our on plaque As and periodontal disease. Elsewhere,cavities suchthe as in a heartor on lungs valve, presence their be life-threat may are Biofilms evenening. moremarine environ in ubiquitous wherements, colonize every they living and nonliving sub strate, everygrain, sediment algal frond, spine, sea urchin’s whale’s back and congregatewaterthe in suspended coating as well, column, particlesdetritus formto “ gregarious nature,are and biofilms by a prevailinglifestyle. chemical Many are signals to orchestrate used bio in life produce Bacteria continuously and detect molecules films. stromatolites recent and found, stromatolitesto form,continue ocean the in Microbial also occur as patches mats ofand scum today. floatingslime, waterin rocks wet and coating bodies and sed deep the in intertidal ocean or on iments, at cold seepsflats, marshes salt and lagoons, and gassprings hot in hydrates, and mangrove swamps, and even lakes and ponds, puddles. These unprepossessingpatches of scum, called areimportant becoming of subjects research. scientific areBiofilms microbial to surfaces. adhering communities Theseliving veneers damp surface,aboutcoat just biotic any or abiotic,evenplanet. Some biofilms the on sun- occur on baked desertrocks, creating"desertcalled coating a colorful varnish."areBiofilms composed of a variety of microorgan whichisms, togethersecrete and them matrix, glueing a gooey substrate.to the them fixing microscopic Like biofilms cities, ------chlo Endosymbiont , power the cell's . They grewthickness in mitochondria . Primitive microbes congregated together, . Primitive microbes congregated together, . Othersthroat, nose, mouth, inhabitour eyes, ecological succession 2 , whereoccurs. photosynthesis The cells ingested (C&H, p. 73). Aerobicoceans ancient the bacteria in (C&H, p. of(about skin areathe of a postageon the stamp) 2 accumulated below, building into calcareousinto building called columns below, accumulated within. Thesewithin. changed time, muchcommunities over ancient as a foreststructuredis and develops time, through over the process of as livingmicrobes upward multiplied and layersof debris formingpatches of grewthat scum availablethe all cover to Organismsseas. ancient the shallow space in differentwith capabilitiesand requirements took up residencedifferent in layers of Primitivebacte photosynthetic communities. these ria grew near the surface and various heterotrophs resided The Scum of the Earth the ScumThe of yearsleast 3.5 At billion long before the first ago, eukaryotes appeared, microbes began formingas associations intimate microbial mats were gradually transformed from preyendosymbionts, into permanentinto finally and structures of Thecells. host the nucleusand other membrane-bound organellesof eukaryote arecells arisen throughto have thought processes. similar were engulfed by anaerobicwere prokaryotes engulfed by and transformed evolutionaryover time into whereplants cyanobacteria respirationplace. Similarly, takes other prokaryoteengulfed by became eventually cells roplasts known to us, must extend throughout extend must the biological to us, known world. fact,In eukaryote organismsarose as a resultof symbio accordingsis, generallythe to accepted Theory of bacteria, includinghundreds of livingsymbio in types, of Many eachwith sis us. as with other as well these invis are guests ible to our normal essential functioning. healthy un mostly associations, and complex intimate Equally cm averagecarries adult 1,000 to 10,000 bacteria, except the armpitand groin, which are home to aboutbacte a million ria per cm and urinary tract.are colon human the in Residing trillions isms. We are largely ignorant We ofisms. these relationships, and most of what we do know comes from terrestrial studying arewalking eco We humans. especially and animals, plants ofsystems and bacteria, mites and mutualistic commensal creatures,other tiny Each our pathogens. mention to not 60 BIOL F150: Introduction to marine biology in the forest soil sample. The prokaryote diversity detected diversity prokaryote soil sample. The in the forest those matched which types, 11,400 almost none of about of marinesediment, they foundbillion individuals, 3.1 sample of In a similar organism. types of different 6,000 senting about individuals, repre they tea bag), found billion 4.8 prokaryote a thecontents of soil (about forest of sediment. In 1 gram soil and marine forest sequenced DNA in small samples of in the 1990s researchers Norwegian thisapply technique, to Among thefirst without cultivating. nature, from directly by sequencing genetic material in samples taken diversity of diversity. insights into microbial tremendous profile a It gives ing Metagenomics been described. than has sofar greater is huge,far diversity alone suggests that microbial host species. This particular them unique to a host, many of with each symbiont living typesof of hundreds perhaps organism, bigger in every on and lives both helpful and symbiotic harmful, microbes, of in anywe look, able to live environment. An entourage everywhere present are They evolution. of billion years 3.8 of lifewith a history of tous,flexible andtenacious forms Clearly, incredibly abundant, amazingly they ubiqui are how many twigs on those branches? there, bacteria are and archaea types of But how many different eukaryotes. the from other asthey are each life,asdistant from of tree ontheuniversal main branches thethree fortwo of account We andarchaea, bacteria know that prokaryotes, Microbial Diversity reefs. coral damaged of help in the recovery or culture, tibiotics or antifouling agents, allow us to enhance aqua an may us with new these chemicals provide ing. Someof signals, theseinducing, chemical competinganddefend of phosis into juvenile polyps. oceanmay beacacophony The andtheir metamor larvae coral induce the settlement of for example, corals, Biofilms coating reef these larvae. of that induce both the their life history. cues chemical produce Biofilm organisms the benthic phase of settle onto biofilms to begin worms, andpolychaete seaurchins, corals, including invertebrates, marine of a wide range of and the larvae isms. Plant spores otherorgan of and growth forthe attachment substrates ,” is a recently developed technique that has produced that has produced technique developed ,” is a recently , or“ settlement whole genomewhole shotgun sequenc andthe metamorphosis ------

resent a fourth domain of life! (C&H p. life! "Eye93: on Science") domainof fourth a resent sounusual that theysequences analzyed may rep are even the all species. Some of from discovered been previously four times genes,as many as had nearly photoreceptor of genespublic forms codingfor including databases, new 782 in genes,new a significant addition to thenumbercurrently DNA 1,214,207 species. in these samples included The new and possiblyat least 1,800, detected asmany as50,000, assumedtobelow, was they occur andspeciesdiversity levels low nutrient very Sea,where six sites in the Sargasso from and collectingglobe samples. metagenomic In initial samples the circumnavigating began ics.these researchers In 2003, Venter, metagenom themostambitious use of is perhaps The than that describedby scientists during years. the past 250 times greater in these two small samples alone is several marine microbes -“unveilingmarine microbes the ocean’s hidden majority.” versity, anddiscover the unknown diversity, andexplore of ( Marine Microbes Census of spawned the International 2003 (C&H p.in 216), efforts conservation and target biodiversity MarineLife( sus of taxonomy.” holeof sity as “the black Cen decade-long The diver O. to prokaryote Edward nent biologist, Wilson, refers we don’t know and mayknow, never emi but it’salot!The it to say, Suffice So, there? speciesare how many microbial scientific challenges. thegreatest is one of croorganisms mi of diversity the metabolic your textbook.Understanding outlined in Table are by evolved of 5.1 marineprokaryotes pathways the metabolic made compounds as well. Some of and most human- compound on Earth natural using every of capable are pathways. metabolic nents, Prokaryotes vianew obtain nutrients and energy,and synthesize compo needed to physical chemicals conditions, exploitder different different continually tofunction un They evolve capabilities. metabolic new under anyto evolve their ability conditions, stems from everywhere, tolive microbes flexibility.abolic of ability The but met in their incredible (their morphology) physical forms their of lies not in the variety microorganisms of diversity The ICOMM Global Ocean Sampling ExpeditionOcean Sampling Global ). The goals of ICOMM are to catalog the known di to catalog are ICOMM goals of ). The COML ) begun in 2000 to assess marine in 2000 ) begun , led by Craig , ledby Craig ------61 Lesson 5: Microorganisms And The Tree of Life

------ox , but , but . Their ozone global warmglobal carbon cyclecarbon in our atmosphere, in 2 and , to the atmosphere., to the , the study ofstudy , the effects the applied biologyapplied methane , astrobiology , on the other hand, areother hand, the , on adding an greenhouseeffect taken up by marine microbes photosyn during up by taken 2 greenhouse gas in our atmosphereour in breathe,we that the and Methanogens . ofand distribution origins, livingspace,in and the possible evolutionof extraterrestriallife. Microbes also are valuable ofsubjects areasmany in of study amazing metabolic diversityprovides a virtually unlimited source ofproducts useful potentially Already, and processes. degrade arethey being used to treat pollutants, wastewater, diagnoseand to produce diseases, pharmaceuticalsand most microbial diversity With products. other industrial many in the stratospherein which protectsfrom us le the sun's radiation.thal Marinemicrobes were,fact, in source the of life. Theall formsonly of of most for life they history, its gave Microbialother livingto all rise continues life things. fabricto hold together the of life, as producers, consum ers, nitrogen-fixers and decomposers. They liveand on in organisms,all and necessaryintimate in and associations, They evenas pathogens. formed the organellesof cells. our microorganisms study our attempt to understandin We the oforigins in other basic questions life, and to answer many are they ofsubjects pathogens, the As biomedical biology. research cause and treatmentthe into of even disease. We whento them look ofpossibility pondering the on other life live Some extremophilesthat environments in today planets. Earthon to those be similar may when began,life or similar on other worlds,to those Marslike moon, Eu or Jupiter’s ropa. The of study these organismshas helped to launch a newfield of science: 4. The CO crucialis thesis marine to the only not absorption the for also of CO excess moderatingthe ing other archaea,Mostly methane-producersthese liveanoxic in swamps,tracts intestinal environments, the marine of ani and sewagemals, treatmentwhere plants are they used to breakwaste. down By generating methane, methanogens areproblem the to of contributing global warming, they but harnessedbe also might to produce energy. for methane scale, marine microbesOn a geologic time also are respon The Earth’sactivity for of sible habitability. photosynthetic organismsmarine of billions over years produced the ygen , , ------of of all primary biogeo are them , too small to , too small producers of dead organ secondary con oil and gas de nitrogen fixation decomposers primary consumers ” (C&H pp. 349–350). ” (C&H pp. zooplankton detritus (dissolved organic (dissolved matter), bacteria, archaea and pro phytoplankton DOM dead microorganisms, the on conducting 98% ofconducting marine microbial loop , which we learned about2 Lessons and in heterotrophic , as we will see in the next lesson, and supply and lesson, next the see in will we , as undecomposed autotrophs - whichturnin fall prey to larger marine animals. . In ancient oceans, over millions of millions over oceans, ancient . In years,dead graduallyand "pressure-cooked," accumulated formingthe matured and gasoil depositson which we are so dependent. roles vital Marine microbesglobal in play also chemical cycles Countless other hand, are responsible current for posits microbesdeep-sea became rapidly sedi anoxic in buried wherements, or stopped. slowed Theyis decomposition the marine food web. Thispartcritical web. food marine the of food marine the web“ called the is marine the microbial loop and decomposition by Without quicklymicrobes,a halt. ocean would to grind the in life isms, as muchisms, as half of the organic matter manufactured by marine producers becomes organisms.and excretedleaked, spilled ocean by the into Microbes areDOM and returning consuming responsiblefor whichnutrients, its restto the be lost, otherwise would of dead organicrole the vital playing marine systems, matter in ofreturning the to material this in contained nutrients the where environment, again they become available producto ers.particulatethe to addition In be eaten by anyone else. These anyone be eaten by predatoryeaten by selves - zooplankton sumers Microorganismsare principal also the life. Microbes also are responsible for crucialthis supplying formuseable in nutrient to producers. Microbesof bulk the up make the as well, the graze that tozoans, tiny on whelming abundance and diversity, they support they whelming and diversity, abundance eco marine at nearlysystems every level. They are the sea, the production energythe ing organic and marine other all for materials Mighty Microbes: The Importance of of Importance The Mighty Microbes: Microbes Marine Theimportance of microorganisms though ocean, the in poorly understood,be overstated. cannot over their In 62 BIOL F150: Introduction to marine biology evolving to evade viral diseases, and viruses can “shuffle” can “shuffle” diseases,and viruses viral to evade evolving diversity. microbial in influencing constantlyrole are Microbes toplay also appear an important They and global climate. budgetthe globalcarbon ecosystems,and may affect even nutrients in marine of andcycling energy of the flow fect they life. Byinfecting marine microbes, may profoundly af infect bacteria (as and 100:1, or cells 10:1 they outnumbermicrobial where in the ocean, without a doubt important are viruses debate, self-assembly. their status in the living/nonliving Whatever themselves by (their name means “poison”), replicating organisms and infectthecells of ment in a limited fashion can,however,on their own.They tothe environ respond or reproduce shell, and cannot metabolize rial in a protein genetic mate consisting only of not a cell have membrane, lifethey might belong. do They of onthe tree less where things, as living much viruses to classify whether debated Finally, anote about “ isms through the potential for exploiting yet discovered, microorgan tobe bioprospecting bacteriophages viruses and ” isessentially boundless. fungi ) and all other forms of of ) and all other forms : It has long been - - - - health of terrestrial plants, and were crucial to their initial crucial plants,and were terrestrial health of vital to the are mutualistic relationships most plants.These of intimately they associatedwith the roots are ties,where lichens creating andalgae, istic associations with cyanobacteria in mutual engage others whereas parasites, fungi are of in sediments. Some types rine environments, particularly in other ma as decomposers important probably are They organisms. decomposer the most important they are where forests, bestknown in mangrove are They ied organisms. little-stud (Fig. mostly microscopic, 11).Marinefungi are totheanimal kingdom to plants,the sister group but are Surprisingly,organisms. related not at all closely they are living is no question that fungi, ontheotherhand,are There unknown largely to us. anotherworld are host cells.Viruses genetic material between transferring microbes, the genes of colonization of the land,aswewill see in the next lesson. colonization of . Fungi are better-studied in terrestrial communi better-studied in terrestrial . Fungi are - - - - 63 Lesson 5: Microorganisms And The Tree of Life - Symbiodinium snow methanogen and phylogeny taxonomy morphology shotgun molecular sequencing, whole genome sequencing (metagenomics) epifluorescence microscope Global Biodiversity Assessment, Global Ocean Sampling Expedition, COML, ICOMM mass extinction symbiosis: commensalism, mutualism and mutualism symbiosis: commensalism, parasitism facultative symbiosis symbiosis and obligate endosymbiont ectosymbiont and Endosymbiont Theory mitochondria and chloroplasts zooxanthellae, biofilm, autoin microbial stromatolite, mat, ducer and quorum sensing DOM, detritus and marine microbial loop, • • • • • • • • • • • • • • • - -

- glossary terms glossary Synechococcus Prochlorococcus epiphytes and endophytes thermophile, endolith, extremophile: hyper thermophile, Strain 121, psychrophile, baro alkaliphile acidophile, halophile, phile, and ciliate frustule and test and foraooze, diatomaceous siliceous ooze, miniferan ooze cyanobacteria, picoplankton, and silicoflagellate, algae: dinoflagellate, diatom, coccolithophorid and cryptophyte protozoan: foraminiferan (foram), radiolarian fungi and lichen zooplankton phytoplankton and picoplankton photosynthesis and chemosynthesis Bacteria, Archaea and Eukarya protists, Protista virus and bacteriophage • • • • • • • • • • • • • • your textbook. your Listed below areListed below terms some key Do a self-check this lesson. for understanding. for If you're unclear about the meaning of ofany these terms, re-read in this course the material and check textbook, guide and in your the glossary at the back of 64 BIOL F150: Introduction to marine biology online at eLearning.uaf.edu and in the course materialsyouonline ateLearning.uaf.edu received. andinthecourse a Include thesubject.) hasnoknowledge of someone who something,"describe" doso-don't just list. (To explaining that you to imagine are andshow your beclear understanding, own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 5. 4. 3. 2. 1. Name and describe three types of extremophile, including their preferred habitats in themarineenvironment. habitats theirpreferred including extremophile, types of Name anddescribe three ontheseafloor, abundances with microbial environments? (subsurface) andwithintheseafloor 10 seawater, addinguptoatotalof milliliterof each Synechococcus tiny picoplankton The environment isenormous. inthepelagic microbes of abundance The answer. for your twodetailedreasons Give animalsandbacteriainnature? of diversity therelative of reflection an accurate thousandknownbacteria.Doyou thinkthisis toafew thanamillionanimalshavebeenidentified,compared More flagellates. coccolithophoridsandsilico diatoms, dinoflagellates, foraminiferans, the"skeletons"(shells)of microbes. Compare marine of theskeletalremains withbiogenous sedimentscomposedof covered isthickly theoceanfloor Most of consumers. mostly heterotrophs, are Protozoans thetic producers. mostly photosyn autotrophs, are Algae single-celledeukaryotes. -bothprotists, andprotozoans unicellular algae willoftenhelpyou todothis. Fortwo terms example: the Bebrief, Comparing/contrasting terms. alltheessentialinformation. but alsoinclude of Define thefollowing pairs h. g. f. e. d. c. b. a. cover sheet cover

, not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , DOM andICOMM 121andstromatolite Strain andbacteriophage cyanobacteria ectosymbiont andendosymbiont loop matsandthemicrobial microbial andmolecularsequencing morphology Pfiesteria andBacteria Archaea Written Assignment for 5 Written Lesson Assignment and submit the written assignment to UAF eLearning & Distance Education according to directions found to directions & Distance Education according andsubmit the written assignment to UAF eLearning and may be the most numerous organisms on the planet. There are 50,000-500,000 prokaryote cellsin 50,000-500,000prokaryote are ontheplanet.There may organisms bethemostnumerous Prochlorococcus (Please make your completeandconcise) answers • 29

brief organisms in the pelagic environment. How does this compare environment. Howdoesthiscompare inthepelagic organisms and clear , but, also complete . For example, if asked to asked For. example, if Prochlorococcus and in yourin - - 65 Lesson 5: Microorganisms And The Tree of Life Name two microbialName two relationships and describe their in marine communities mutualists with their hosts. Explain three that microorganisms critical functions perform in marine ecosystems. areHow common Describe three to find them? Where biofilms in marine environments? expect that ways you might chemical signals operate in biofilms. 6. 7. 8. 66 BIOL F150: Introduction to marine biology 67 Lesson 6: The Producers: Marine Autotrophs and Primary Production . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). and "The No-Zone"), pp. 334-335 ("The Phytoplankton"), p. 336 (Table 15.1), pp. 15.1), pp. 336 (Table 334-335 ("The p. Phytoplankton"), and "The No-Zone"), pp. Tides and Harmful338-339 ("Red Algal 221-229 ("The Blooms) and review pp. ofFlow Energy and Materials") Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Understand diversity how the types, and history of marine primary producers differ fromsignificantly those on land. Be familiar with the major types of marine autotrophs. Understand the role of primary producers in marine ecosystems. Understand the factors controlling marine primary productivity. the importanceRecognize of marine primary productivity to Earth's habitability— past, present and future. the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education. C&H: Chapter 6 (pp. 102–113) and pp. 237–239 ("Overwhelming the Nitrogen Cycle" 102–113) and pp. C&H: Chapter 6 (pp. More information, including interactive flashcards, quizzes, videoclips can be and links, 1. 2. 3. 4. 5. 1. 2. 3. Reading assignment Reading steps for completing the lesson the completing steps for This lesson concerns the organisms at the base of the trophic pyramid, the producers that convert the energy supplying and materials inorganic materials into organic matter, necessary all other forms for of life. this Lesson of Objectives Marine Autotrophs Marine Autotrophs and Primary Production The Producers:The 6 68 BIOL F150: Introduction to marine biology T ply us with food,clothing, shelter, fuel,medicines and other to human existence, they sup on land.Crucial production the primary environments and conduct most of most extreme in all but landplants. live the They majorityof prising the vast explosively, andspread Flowering plants diversified now com than 200mya. flowering plants dinosaurs. the dominatedduring the of age like our pines and spruces, mya. 400 about peared environments. more into many andto spread forms, larger much them to evolve later. allowed million years specialized structures These 20–40 andotherspecialized tissues, appeared structures 6.1). (C&H Table other photosynthetic organisms guish them from that distin characteristics biochemical still share two groups andthese ago), mya 460 (million years around algae green 7,000 and about species of 300,000 about plants include Green out by and obvious the familiar tion is carried produc primary thevital function of world, In our terrestrial Seeding the Land in perspective. and in the sea, tohelp put these differences onland producers of theevolution of history gin with a brief environments. We be in marine and terrestrial greatly differs called process, life. This of other forms matter, forall andmaterialsnecessary supplying theenergy Vascular plants

ducers the pyramid, trophic the at the base of the organisms his lesson concerns Angiosperms ) that convert inorganic materials into organic materials into organic inorganic ) that convert green algae green , were the last major group toappear, the last major group , were less Seed-bearing , land plants with water-conducting Gymnosperms (seeds enclosed in an ovary), or in an ovary), (seedsenclosed producers . Land plants evolved from from . Landplants evolved (Marine Autotrophs and Primary Production) andPrimary (Marine Autotrophs primary production primary vascular plants ap vascular (or ("nakedseeds"), The producers The green plants green primary pro primary land plants • ------, .

have already destroyed more than 50% of their coverage. of than 50% destroyedmore have already year,per 1% of and at arate rainforests destroying tropical bydescribed animal species sofar science.We currently are all plant and of than half contain more which rainforests, in tropical occurs and productivity plant diversity flowering oxygen peak of and dioxidecarbon The in the atmosphere. is vital to the balance of productivity useful chemicals. Their the ocean swarmed with simple prokaryote cells. with simple prokaryote swarmed the ocean life’shistory, of twobillion years For undiscovered. the first many still organisms, microscopic in the ocean are ducers pro majority of the vast counterparts, Unlike their terrestrial ton Microbial Producers: The Phytoplank for requirements greater oxygenwith multicellular organisms, alsoallowedlarger up of build- by (also damaged This human activities). stratosphere layera protective of created which mosphere, added bacteria andarchaea photosynthetic activity, of marine years billions of Through by bombarded the uninhabitable, sun’s was lethal radiation. history,the land Earth’s forthem. For habitable most of environments for making terrestrial responsible also are andanimal life onland,marineorganisms the plant,fungal Besides seeding phyla evolved. hadalready ly all the modern near where the sea, alsocamefrom andvertebrates, brates land animals, both inverte first most plants. The health of tothe manymutualistic became symbionts, now necessary risetoland plants.gave onland and eventually emerged algae green the first before than 3 billion years for flourished more by life in the which sea, possible made on land was and diversity All this productivity Fungi aerobic respiration aerobic oxygen emerged with the algae and with the algae emerged tothe ocean andat ozone , to evolve. to evolve. , Micro in the ------69 Lesson 6: The Producers: Marine Autotrophs and Primary Production

, ------hy sym Prochloron dinoflagel , primarily in , primarily , microbial chemosynthetic Hydrothermal vents have been found sup have been found free-living , a genus of , a genus , whereand sulfide hydrogen , ammonium and ironand , ammonium synthe to , livingof tissues the within of many symbionts conduct photosynthesis, photosynthesis, conduct symbionts free-living plankton methane cold seeps cold Symbiodinium , . chemoautotrophs endosymbionts also contribute to primary contribute also production sea. the in , are zooxanthellae who, are partner zooxanthellae corals with and are like whaleslike which also produce and meth sulfide hydrogen chemoautotrophsane. Many are Oth or growingsediments thickinto mats on the ocean floor. ers are which animals the inhabitrich,these environments. lightless Most primary productionthe ocean, fueled in as on land, is Thedark, deep-sea cold was environment solar energy. by to be and unproductive.lifeless long thought Organic matter epipelagicdown to lower sinks produced phytoplankton by meager fare is it but levels, vastthis in realm. known now is It some bacteria that and archaea thrivethese environments, in do everywhereas they else. Innovative prokaryotesenergythe use chemicalsin contained like drogen sulfide certain In organicrecentsize until unknown habitats, matter. these decades, porting richincredibly communities. the deep-seamid-ocean ridges associated with springs hot and trenches, vare surrounded flourishingofcommunities by tubeworms, clams, other invertebrates,and mussels fueled chemosynthetic bacteria and archaea.by communi Similar at also exist ties even and at deep-sea seepmethane from out sea floor, the “graves,”remainsdecomposing the of large dead animals gos are fur the and gos ingest, cyanobacteria the colored they by of polar bears cyanobacteria.green sometimes tinged is by MicrobialOther Producers the to addition In bionts Photoautotrophic providingenergy and organicre in hosts, to their matter turn Theother benefits. a home and are best known for the zooxanthellae lates formationto the critical of coralreefs. form Other symbionts foraminand even tiny sponges mollusks, with associations ifera.bacteria of Photosynthetic genus the provideservicessimilar sea squirts. the hosts, their to , , , ------and , and , which photo Prochlo diatoms , and many , and many . About the . About chemosyn Endosymbi brown algae and brown,red algae

, microscopicphoto , first green arose as a resultof cells , as well as , as well ) have revealed) most the green bacteria , easily caught in standardin caught , easily . The firstpro multicellular and algae and Synechococcus seaweeds , too small to be caught in nets, nets, in to be , caught too small red . Eukaryotic algae include sulfur-oxidizing bacteria sulfur-oxidizing and pathways for carrying for and pathways out photo phytoplankton purple , as unicellular as well cyanobacteria , and some photosynthetic bacteria, pri , and some photosynthetic chloroplasts archaea dinoflagellates . More previouslyalgal minute cells, unknown, , single-celled picoplankton . Thesecyanobacteriaminuscule are members communities developed, communities including photosynthetic and molecular sequencing, metagenomics sequencing, molecular forms such as Theory cryptophytes

The Red Sea gets its color its fromThe Sea gets red Red cyanobacteria, flaminpink synthetic pigments 5.1, 5.2 and 6.1). Cyanobacteria are Tables (C&H synthesis particularly diverse,an assortment with of photosynthetic impartingpigments a rainbow of colorsorganisms.to these green algae 92: also are being discoveredmetagenomics (C&H p. using Big Surprises"). Cells, "Tiny The major groups of phytoplank 336). 15.1, p. (Table areton textbook your summarized in Thesegroups have evolved variouslight-capturing otic members,otic includeabundant,of types most the many cyanobacteria, plus photosynthetic silicoflagellates coccolithophorids, dinoflagellates, and rococcus of tiny the and are arguablyorganismsthe most abundant on Earth. The diverseare phytoplankton Prokary a hugely group. marily cyanobacteria (formerlymarily called “blue-green algae”). as we learnedpreviousthe in newlesson, However, tech ( niques epifluorescencemicroscopy phytoplankton: abundant still carriedstill by out organismssynthetic waters. sunlit drifting in Phytoplankton to include been known have long algaeunicellular suchas diatoms nets plankton followed by multicellular by followed firstthe same time, appeared.animals multicellular tiny more95% than of marine primary productionis Today, eukaryoticappeared. cells According to the ont cyanobacteria, like cells, photosynthetic smaller engulfing which became ducers were leaf-like tiny bial mat organismslike thetic derive energy molecules, fromsulfide hydrogen splitting rather fromthan Nearly sunlight. years two billion later, 70 BIOL F150: Introduction to marine biology presence of water and nutrients in the soil, have shaped the and nutrients in the soil,haveshaped water of presence air, andthe tothe ground, penetrates light readily in which lowdensity of environments.The different ing invery two for adaptations liv of the result and those on land are ers produc betweenmost marineprimary differences stark The DifferentTwo Worlds trollably, disease. speciesand spreading destroying native can upsetdelicately uncon balancedecosystems,growing distances. species Non-native over great to port, port from ships traveling of ried on the hulls or in the ballast water of is the introduction threat increasing ment. Another rapidly caused bypollution problems and other coastaldevelop and sources, freshwater of and filling, diversion dredging of result asa dramatically in many placeshavedeclined and othercoastal habitats beds Seagrass ignored. stood or under often has been poorly and erosion, flooding tion of for the preven fish, and many other animals, andas buffers spawning birds, formigrating ashabitats importance Their by threatened many human activities. are they support, animal plants,communities marine flowering andthe rich and home toseaweeds coastalenvironments that are The bays andtropical and estuaries. in temperate widespread es pickleweed of and shrubs thetrees plants include Marineflowering onEarth. ductive they dooccur,where the most pro however, someof are habitats toshallow coastal environments.The restricted are plants ing systems,few toterrestrial In distinct 6.1). contrast physiologically land plants (C&H Table distinct from and are leaves, stems and roots, to true land plants, seaweedslack habitats. Although similar they other shallow-water appear the in some coastal habitats. Multicellular algae, producers important are picoplankton, multicellular organisms ered discov recently are which out by phytoplankton, most of is carried production marineprimary of While at least95% Multicellular Producers (more closely related to lilies than grasses) which are are which tolilies than grasses) related closely (more invasive seaweeds are marine producers. Like the seaweeds, they marine producers. are

found in saltmarshes, andthe found in saltmarshes, species mangrove forests mangrove , are common along rocky shorelines and shorelines commonalongrocky , are (both plant and animal), often car , the cordgrasses seagrass flower and ------through their cell walls, but are restricted to shallow habi restricted their butcell walls, are through and nutrients directly water seaweedsalso absorb larger The shells bearing spines and pores. with elaborate drag, and their area, their surface cells like diatoms increase tio their weight and maximizedtheir tively high frictional buoyancy ontheir low weight and rela is dependent Their is enough light forphotosynthesis to occur. there where suspended in the photic zone, also can remain walls. They their cell through and nutrients directly water can absorb phytoplankton situation. Microscopic different ates avery ily cre accessible, but as it penetrates, light decays rapidly seawater, read anddissolved nutrients are water in which sunlight and carbondioxide. high density of to absorb The tothe sky and leavesraised and nutrients, andtrunks water totakeup in the ground land plants. haveroots large They tion also is more difficult to measure in marine systems. to measure difficult tion also is more produc to detectin the ocean, soprimary difficult more much are producers the systemovertime. Just as primary in changes health and useful fortracking ing its overall in the system, indicat is happening sign” telling us what an ecosystem is like taking its pulse: it is a key “vital of productivity Measuringthe primary and globalclimate. the globalcarboncycle mans, influences andstrongly for hu resources important are which ganisms, many of all other marine or supports productivity marine primary since in these values, interested extremely Scientists are time,for example: amount of in a given seasurface ter of me carbon fixedunder one square as the amount of time,usually expressed per unit of production primary of The dioxide carbon takenup. the amountof or fixed, carbon usually asthe amount of expressed producers, by primary production Primary Primary Production nutrients. and to absorb to the substrate, to attach roots use their relatives, plants,rine flowering like their terrestrial Ma to the substrate. can attach their holdfasts tats where , giving them enough drag to prevent sinking. Larger toprevent them enough drag , giving primary productivity primary drag grams C/m grams is the amount of is the amount of . The tiniest cells have minimized . The of asystem is the amount of 2 /year surface-to-volume ra surface-to-volume biomass created created ------71 Lesson 6: The Producers: Marine Autotrophs and Primary Production

------). hypoxia Red tides Red nitrogen-fixing occur, whenlight occur, , a massive, increasethe in ,” created eutrophication by . Ironwas sea, ancient the in plentiful spring blooms iron dead zones dead eutrophication . They are a periodic natural occurrence, triggered Pacific, Pacific, and Southernadded iron When scientists to Oceans. dramaticthese environments, resulted. blooms Thisprompt ed controversialproposals to fertilizeareasthese iron, with increaseto both productivitywarming,global slow to and by uptake. Massive increasingstorms dust carbon dioxide are an importantsource of ironlargein areas ofocean. About the yearly blown is metric tons 500 million fromSahara the des ert,Norththe into settling Atlantic, whereprimaryboosts it blooms favorabletemperate In by conditions. environmental and polar waters, annual and warmth return can take advantage and phytoplankton ofthe darkerduring accumulated nutrients months. winter provideThese most of may blooms seasonal primary the production the entirefor regions.these in year (which aren't red always tides), are don't involve and an other type of Theyspecies a single bloom. areby caused rea unknown for ofand occur unpredictably, phytoplankton some species of tides involving Red sons. dinoflagellates, waterscoastal foul and cyanobacteriadiatoms periodically around the world, sometimes producing toxins. deadly They seem to be increasingly frequent, as a result possibly of increased activities human along coasts. and pollution Primary productivitylarge in areas of open ocean are limited micronutrient, the by shortin is today but subarcticthe in equatorial Pacific, supply for producersfor to use. The presence of cyanobacteria, nitrogen-fixingand other and ar bacteria chaea, strongly influencesprimary productionmarine en in Other naturalvironments. sources of fixed nitrogen marine in include environments from input rivers by and electrification are humans the source of However, lightning. of most the re from sewage release, run-active nitrogenthe ocean today, in offof agriculturalfertilizers rivers,into and burningof fossil Thisfuels. degree ofenrichment, nutrient nutrient actually causes pollution, growthof algae. massive Massive de growth by followed is deprivingcomposition, of environment the ( oxygen or “ zones, Hypoxic suchofmouth as at the Mis the places today, many in exist westward extending Gulf along the River, sissippi of Mexico. increasesSudden growththe in of are phytoplankton called

, . ------) photic dysphotic reactive nitrogen limiting nutrients zone, no light penetrates. zone, no light , watersince avail always is aphotic , those required producers by in . At lower light levels, in the in levels, light lower . At , also required, also doses (in smaller in but limiting factors , nitrogen makes up 78% of our atmosphere, 2 macronutrients net productivity but it must be fixed, or converted must it but into flagellates, silicon may be the limiting macronutrient, be the limiting as it is may flagellates, silicon nitrogen is Most often, however, shells. their needed to build and adequate light with environments in nutrient, the limiting warmth.N As productionscale. Organismsa local on vary their in availabil and nutrient nutrients, essential need for other factors.extremely variable,is ity many influencedby In the case of diatoms and silico some algallike species, ily available as dissolved carbon dioxide, but nitrogen, but available ily dissolved carbon dioxide, as phosphorus become may and silicon Micronutrients limit manganesemay and zinc), cludingiron, copper, but often the opposite is true.opposite is often the but Thisbecause nutrient is availabilityalso strongly influencesproduction. Availabil ofity productivitylarge limits often doses, on a regional scale. Carbon, most importantthe macronutrient, readalways is changes in solar radiation, cloud cover, and the claritythe and of changesradiation,solar in cloud cover, Muchof water. the radiationsolar incoming the into goes ratethe Since ofthe water. heating also is photosynthesis dependent on temperature, seem that productivwould it decreasewould ity from steadily poles, the equator to the zone, phytoplankton arezone, phytoplankton photosynthesis not able to conduct at all. Deeper the in still, Organismsdiffer sensitivity their in inten and light to light, and seasonal dependingdaily latitude, on variessity widely, fallsoff to a point diminishing rapidlypenetrates as it water, where the rate ofthe supply to enough only is photosynthesis ofneeds maintenance organism, the and there no growth, is or are the usually environ words,nutrient-deficient other and able. In lightless arements the “deserts”of the sea. The vast majority of marine producers therefore liveepipelagic the in ( relativelyand zone, light-filled warm. Theofintensity light Primary productionofa function is factors, many including temperature,light, availability and the of variousnutrients Primary productivityby limited usually on land is and water. In ma temperaturethat order. in and light, water availability, availabilitynutrient light, environments, rine and temperature 72 BIOL F150: Introduction to marine biology lion square kilometers of ocean, whereas for ship a travel whereas ocean, of kilometers lion square satellite scan might contain 2 million pixels, covering 2 mil productivity. primary A 2-minute of globalmaps real-time, computerized mathematicalscientists models,with provide using nologies tech ships. sampledfrom Newer in seawater chlorophyll, by light absorbed (or oxygen orthe amount of off), given carbondioxide consumed measuring the amount of od of meth to calculate by it's difficult thetraditional vironment, andcomplex en in this vast is so variable Since productivity bothtotimeandlocation. with respect variable tremely and ex powerful, are production onprimary effects Their water, onasmall scale orglobally.cause movements of Many ocean. complex in the tropical processes productivity stratified more waters tendstoleavetropical sonality in the tropics sea of lack tofuelthe spring bloom. The ready the surface, to water months, deeper sinks and displaces nutrient-rich overturn of asaresult waters in polar and temperate Mixing occurs highly productive. are thatthesecoldwaters one reason the equator, Antarctica, Oceanaround and in the Southern alongcoastsandat Upwelling occurs the surface. toward especially by mixing is accomplished byThis component nutrients to the surface. its of onits decomposition andthereturn depends pelagic in the epi production But continued primary sea consumers. andmaterials for deep- energy of source matter is a crucial pump -the theseafloor nutrients toward of net transport in a matter results organic on dead gravity constant pull of in the sea, ason land, causing material to sink. The rules ever, andbottomsediments.Gravity waters is the deep-sea nutrients how in the ocean, of source mostimportant The Pacific, aswell. and Mongolia tothe North contribute iron China northern of originating in the Gobi deserts dust storms blooms. Massive and sometimes causes harmful production we learned about in Lesson 4. Sinking dead organic in Sinking Lesson 4. organic dead about welearned , facilitated by wind, waves, currents and tides, and byfacilitated , currents wind,waves, . Surface water,. Surface denser during made cold winter , resulting in lower but relatively constant primary constant primary in lower but relatively , resulting upwelling remote sensors remote , the slow persistent rising of water water rising of , the slow persistent carried by carried satellites carbon , and ------production (C&H Tableproduction However, 10.1). in Les aswelearned contribute equally about toglobalprimary producers primary by ered the ocean adds up, sothat marine and terrestrial cov environments, area but thevast terrestrial productive than the most less productive Many oceanenvironments are important. ing dynamicsocean particularly has become ocean (and global) systems,understand at a time when a year. our knowledge of is revolutionizing technology This by compositing images over aweek,month or patterns their bloomsused todetect color imagescanbe andtrack tion) as to by referred imagesare phytoplanktonThese concentrations. of chlorophyll-a of detect levels sensors remote The area. just one pixel of tocover years than 11 km/h,iting would at 20 take more portant to our understanding of our rapidly changing world. changing ourrapidly of to ourunderstanding portant critically but im are difficult extremely are trends, long-term and predict that have been occurring to decipher changes in marine environments,and tocalculate productivity Efforts plays production in the global carbon cycle. marine primary that considering the significant role concerning, very they are correct, these estimates are seas. If in polar andtemperate forspring blooms necessary andinhibits the overturn waters ocean of stratification more creates which temperatures, face atmospheric and seasur with increased corresponds This by than globally,6% has declined more 1980. since about productivity NASA scientists calculated that marine primary the sequestration carbon as in ocean water.tain times 25 as much This andaccumulates sediments.in ocean Sediments con posers decom of reach matter in the ocean sinks out of organic dead the of But much to the atmosphere. oxide returns promptly matter decomposeson land, carbon di When dead organic and marineecosystems. betweenterrestrial greatly differs fixedcarbon as in carbon the air.times as much of fate The son 3, CO greenhouse effect greenhouse NASA ocean color 2 is highly soluble in water, andthe ocean holds 50 (National Aeronautics and SpaceAdministra (National Aeronautics near the ocean surface, producing images producing the oceansurface, near has been hugely important for moderating for moderating has beenhugely important (C&H Figs.Ocean and 10.19). 1.13 and global warming . A study by carbon ------73 Lesson 6: The Producers: Marine Autotrophs and Primary Production - - - macronutrient and micronutrient reactivenitrogen-fixation, nitrogen carbon pump/sink/sequestracarbon-fixation, tion greenhouse gas, greenhouse effect global warming, acidification and red spring bloom tide bloom, and dead zone eutrophication, hypoxia invasive species ocean color ozone layer chloroplast (chlorophyllphotosynthetic pigments a, c, carotenoids, phycobilins, etc.) surface-to-volume drag ratio, primary producerand primary production productivity and net productivity photic, dysphotic and photic zones overturn,upwelling, stratification limiting factor/resource • • • • • • • • • • • • • • • • • • - - - - glossary terms glossary Prochloron , Prochlorococcus Symbiodinium and chemosynthesis and photosynthetsis (chemo autotroph and photoautotroph) angiosperms/flowering plants (halophytes: seagrasses, cordgrasses, mangroves) free-living, endosymbiont, ectosymbiont, Endosymbiont Theory zooxanthellae, thallus, stipe, blade, holdfast and pneumato blade, thallus, stipe, cyst sporophyte and gametophyte alternation of generations celled algae) (green, brownseaweeds and red multicellular algae) calcareous greensea lettuce, algae, rock corallineweeds, kelps, algae Synechococcus single-celled algae dinoflagellates, (diatoms, coccolithophorids, cryptosilicoflagellates, phytes, and brown, red and green single- phytoplankton, picoplankton, plankton net phytoplankton, picoplankton, cyanobacteria, purple green bacteria, bacteria bacteria and sulfur-oxidizing • • • • • • • • • • • • • your textbook. your Listed below areListed below terms some key Do a self-check this lesson. for understanding. for If you're unclear about the meaning of ofany these terms, re-read in this course the material and check textbook, guide and in your the glossary at the back of banks area there is no need to complete the form. Come to our office is no need to complete the form. Come to our office there banks area 4 hours for complet during business hours to take your exam.) Allow ing the exam. After completing the written assignment for this lesson you will be After completing the written assignment for this and complete in your area for the first exam. Locate a proctor ready the Exam Request Form on the following pages. (If you live in the Fair 74 BIOL F150: Introduction to marine biology online at eLearning.uaf.edu and in the course materialsyouonline ateLearning.uaf.edu received. andinthecourse a Include thesubject.) hasnoknowledge of someone who something,"describe" doso-don't just list. (To explaining that you to imagine are andshow your beclear understanding, own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 1. 5. 4. 3. 2. Define, by comparing and contrasting (youDefine, by knowthedrill),following comparingandcontrasting terms: tion: photosynthesis, carbon-fixation, decomposition,calcium carbonate,andcarbonsequestration. thefollowing possibilities, inyour toinclude but descrip various besure are There toitthere. happens scribe what intotheocean,andde carbondioxide theatmosphere from Follow intheglobal carboncycle. role amoleculeof productivity, primary but they play larger amuch Earth's of half for about responsible are producers Marine primary tidesandeutrophication. springblooms, red Describe andcompare andlocally. regionally intheocean. production productivity primary primary influences Describehowoneotherfactor affect which clarity, withlatitude, daily cover and water andseasonalfluctuations, cloud intensity alsovaries allof highest.Light are tosynthesis cannotoccur. lightlevels where takesplaceatthesurface, production Mostprimary pho level, by influenced isstrongly light.Whenlightintensitydiminishesbelowacertain productivity Marine primary inyour answer) marineproducers 5specificexamples of (include marineproducers? themostimportant intheocean?Whoare theproduction forplants alsoresponsible mostof flowering plants. flowering onlandisconductedby Are production thehugely diverse theprimary mostof By far f. e. d. c. b. a. k. j. i. h. g. cover sheet cover

, not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , dead zonesandoceancolor andmicronutrient macronutrient sporophyte andgametophyte aandcarotenoids chlorophyll andchemoautotroph photoautotroph andkelpforest mangal Synechococcus andsealettuce silicoflagellate andseaweed seagrass phytoplankton andzooplankton photic zoneanddysphotic zone Written Assignment for 6 Written Lesson Assignment and submit the written assignment to UAF eLearning & Distance Education according to directions found to directions & Distance Education according andsubmit the written assignment to UAF eLearning and (Please make your completeandconcise) answers Symbiodinium • brief and clear , but, also complete . For example, if asked to asked For. example, if in yourin - - - 75 Exam Request/Form - - UAF eLearningUAF (Note: photocopying or taking notes from any examination paper or taking notes from is not permitted); examination any (Note: photocopying UAF eLearningUAF violation of process. the examination verification of instructional materials (if process; allowed during the examination any) return of all papers, including scratch and the completed Examination Cer questions, examination sheets, tificate to termination of confiscation of the examination, and immediate notification of materials, exam telephone ifby there is improper conduct on the part of evidence that there the student or any been a has examination session; examination identification of (or other verification, if photo I.D. the student by necessary); provision of area a quiet, well-lighted as free from noise and distraction and within supervisory as possible distance of the proctor; security of until it is opened in the student’s presence the sealed examination at the beginning of the • • • • • • Please sign the student’s Examination Request Form beforePlease sign the student’s Examination Request s/he returns it to our office and keep this Information referenceLetter for when administering the examination. advance by contacting UAF eLearning contacting UAF friends cannot be and/or advance Relatives by at 907-479-3444 or [email protected]. proctors. Proctor responsibilities include: stand their responsibilities and agree to ensure that integrity. When an individual is accepted as a proctor, s/he When an individual is accepted as a proctor, stand their responsibilitiesand agree to ensure that integrity. represents the University of process. the examination for and is accountable Alaska Fairbanks Proctors or an administratorcommunity college schoolat a public site be education officials must at a university, such other governmental persons or community officials, or if, or library, are unavailable, other people approved in You have been designated as a proctor Independent for Learning examinations. You The integrity of process the examination is fundamental to our program supervised because it provides the only thereforecheck are We of concerned that prospective proctors and capability. the student’s knowledge under INFORMATION LETTER TO BE GIVEN TO YOUR PROCTOR YOUR TO GIVEN BE TO LETTER INFORMATION

77 Exam Request/Form Exam 1

http://elearning.uaf.edu Date: Phone: Phone Examination No:

UY4

Fairbanks, AK 99775-6700 Fairbanks, University of Alaska Fairbanks Section No: 907-479-3444 fax: 907-479-3443 UAF eLearningUAF & Distance Education

2175 University 756700 Avenue South, Suite 200 , PO Box EXAMINATION REQUEST FORM REQUEST EXAMINATION

Introduction to Marine Biology BIOL F150

The exam will be mailed directly to the proctor, so maintain contact with that person. so maintain The will be mailed directly to the proctor, exam request submit an exam from eLearning may the UAF you website: Alternatively, Send this request form in a pre-paid provided with course envelope or to: materials Select an approved testing site and arrange individual an appointment to proctor with an eligible the examination. Ensure the proctor that has read the attached Letter of Information Form. Schedule this Request and has signed farthe exam enough in advance receipt to allow for of preceding all lessons of and the mailing the exam the test itself. If area live there in the Fairbanks you this form. need to complete is no office Come to our during regular business hours exam. to take your 4. 5. 3. 2. 1. Student's Email: Student's signature: Student's Address: Information Letter, and agree to observeInformation Letter, all the requirements of process. the examination Proctor’s signature: I agree before taking it. examination, this to complete all lessons covered by Proctor's Email: I have read and understand the duties and responsibilities of proctor an examination as described in the attached Proctor’s Name & Title: Proctor’s Organization: Mailing Address: Student Name: Course No: Course Title: PLEASE PRINT OR TYPE INSTRUCTIONS

79 Lesson 7: Marine Invertebrates—The lower phyla - Marine

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). Be aware of the basic trends the evolution in of from to chor animals, sponges dates. Be familiar with the "lower" invertebrate plans and life histories. their body phyla, the information of pages Read this lesson in the following for this Course Guide. Complete the Biology Complete the eLearning & Distance Education. Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e 1. 2. 1. 2. 3. 295 ("Life in Mud and Sand") 115–149) and p. C&H: Chapter 7 (pp. More information, including interactive flashcards, quizzes, videoclips can be and links, Reading assignment Reading steps for completing the lesson the completing steps for Moving up the trophic up Moving pyramid, we turn consumers, to the the heterotrophs that rely directly or indirectly on producers their energy for are and organic already We materials. familiar with the smallest consumers, heterotrophic the archaea and bacteria, and the the more Now we beginprotozoans. and familiar to study consumers, visible the members of the animal kingdom. this Lesson of Objectives Invertebrates— lower The phyla Marine 7 80 BIOL F150: Introduction to marine biology Subkingdom Eumetazoa Subkingdom Parazoa Kingdom Metazoa Domain Eukarya Bilateria Radiata Superphylum Lophotrochozoa Superphylum Platyzoa Superphylum Lophophorates Trochozoans Phylum Cnidaria Phylum Porifera: sponges Phylum (goblet Entoprocta: worms) animals) Phylum Bryozoa/Ectoprocta(bryozoans/moss worms) Phylum Phoronida(horseshoe (lampshells) Phylum Brachiopoda Phylum (spoonworms) Echiuria: Phylum Sipuncula(peanutworms) worms) (ribbon Phylum Nemertea Phylum Annelida Phylum Mollusca Phylum Cycliophora Phylum Gnathostomulida Phylum Rotifera Phylum Gastrotricha Phylum Platyhelminthes(flatworms) combjellies Phylum Ctenophora: Class Hirudinea: leeches Class Hirudinea: etc. earthworms, Class Oligochaeta: Class Polychaeta: marinesegmented tuskshells Class Scaphopoda: Class Polyplacophora: chitons octopuses, squids,Class Cephalopoda: etc. mussels, clams, oysters, Class Bivalvia: snails, limpets,Class Gastropoda: abalo Class Cestoda:tapeworms Class Trematoda: flukes Class Turbellaria: flatworms freeliving Class Cubozoa:cubicaljellyfishes sea Class Anthozoa:seaanemones, corals, jellyfishes larger Class Scyphozoa: colonies bushy/feathery Class Hydrozoa: worms cuttlefishes, etc. etc. nes, nudibranchs, fans, etc. minutemedusae polyps, release of Fig. 12 Animal ClassificationFig. 12Animal - Superphylum Ecdysozoa Superphylum Superphylum Deuterostomia Superphylum Phylum Arthropoda Phylum Chordata worms) (acorn Phylum Hemichordata Phylum Echinodermata worms) Phylum (arrow Chaetognatha Phylum Loricifera Phylum Kinorhyncha Phylum Nematoda(roundworms) Subphylum Chelicerata Subphylum Vertebrata Subphylum Vertebrata Subphylum Cephalochordata Subphylum Urochordata Subphylum Uniramia Subphylum Crustacea reptiles, birds, mammals) birds, reptiles, phioxus) Class Thaliacea: salps Class Thaliacea: larvaceans Class Larvacea: Class Ascidiacea:seasquirts Class Crinoidea:sealillies, featherstars sea cucumbers Class Holothuroidea: seaurchins Class Echinoidea: brittlestars Class Ophiuroidea: seastars Class Asteroidea: Class Chilopoda:centipedes Class Diplopoda:millipedes Class Insecta:insects shrimp, crabs, lob Class Malacostraca: copepods, Class Maxillopoda:barnacles, shrimp, water fairy Class Branchiopoda: mites ticks, spiders, Class Arachnida: Class Pycnogonida: seaspiders crabs horseshoe Class Merostomata: sters, krill,amphipods, isopods sters, ostracods etc. fleas, (fishes, amphibians, (tunicates) (lancelets–am - - 81 Lesson 7: Marine Invertebrates—The lower phyla , ------), verte is one is inverte , this great, this chordates ( ) and ) Cambrian Period backbone Cambrian Explosion Subphylum Vertebrata , or a Phylum Chordata . phylum vertebrae (with a backbone). (without Thisa biased view, quite is About 550 mya to 500 mya, during the during to 500 mya, 550 mya About bursta huge of evolutionary innovation took place the an in as the ocean. cient Known proliferationof life produced most ofpresent-day the animal of explosion An phyla. eukaryotic life was in made possible partactivity the of by prokaryotes,marine photosynthetic hundredsduring ofof millions years,oxygen adding enough to support environment to the greatthe metabolicneeds of fied. The other 2 chordate the other animal all plus subphyla, arephyla, invertebrates.devote will to the in two lessons We , surveyingabout half of (a cur 35–40 the phyla sorydiverse and introduction huge this to group). This lesson describes some ofmore the primitiveinvertebrate In phyla. we survey lesson, next the more the advanced invertebrate 9–11, Lessons vertebrates. and, in the phyla mate Use the Course this rial in as an introduction Guide to these groups, which are Re described greaterin textbook. your detail in learned whatcalling you 4, each Lesson consider how in of groups successful these highly evolved has adapta unique requirementsbasic the to fulfill tions of energy life - obtaining and raw responding materials, externalto the environment growinghomeostasis, and maintaining and reproducing. the invertebrateAll have marine representatives, phyla and some are exclusivelyfact, marine. In most ofcurrent the ani some others, (plus mal phyla were now extinct) presentthe in sea more before life appeared500 than mya, any on land. Traditionally, animals are animals dividedtwo groups: into Traditionally, brates brates 97% at least ofthat considering species are animal inverte brates. Vertebratesa group one subphylum, just constitute a largerwithin of three within subphyla whichone of is just currently the 35–40 phyla animal identi . ------• het shown Metazoa ofmeta the , or Tree ofTree Life relationshipsmajor among the Kingdom Animalia in the Appendixthe in of Course this Guide. ) are multicellular, eukaryotichetero are) multicellular, taxonomic classification taxonomic

Marine Invertebrates—the lower phyla lower Marine Invertebrates—the phylogenetic who rely directly or indirectly on the produc metazoans

conspicuous, but by no means least significant marine significant least means no by but conspicuous, organisms- the diverse marine microbes and produc n the previousthe n learned we lessons, aboutleast the sent just one twig on the eukaryote on the one twig just sent branch, and eukaryotes one of just for account three the branches main of life. For a geologic perspective on the appearance of the different groupsanimal general)in life (and Earth,on the at look Geologic Time Scale their characteristics.their These are organizingtools for useful and reviewing greatthe diversityof Keepmind in animals. on the occupy animals that position the Figurein 11, and picturedfirstthe on page of eachchapter repre animals great Despite their textbook. diversity, your in ring backring figure to this picture""big the to get ofclas animal referAlso sification. which to Figure textbook, 7.54your in the shows Therephyla. animal end ofat the also a table is eachtext coveredanimals the listing and summarizing book chapter, the most diverse.the This reflectmay diverse the they ways have evolved and to avoid becoming another food to obtain to manufactureunable meal, being animal’s food. own their Figurea 12 shows different the study you As groups,animal keepzoans. refer Animals ( Animals Of ingestion. trophs by food obtain that the 3 ofkingdoms eukaryotesareanimals multicellular fungi), animals, (plants, ersenergytheir all have for and organic We materials. alreadyconsumers, smallest met the heterotrophic the archaea and bacteria, and the protozoans. Now we are ready to meet the more and familiarconsum visible ers,members the of ersof (most whom are microbial).food the up Moving chainfrom producers,consumers, we meet the the erotrophs I 82 BIOL F150: Introduction to marine biology are are the Subkingdom Eumetazoa of members most primitive The Radiata these materials( nation of spongin ( the protein of fibers ( calcium carbonate including support, rial usedforstructural mate tothetypeof according classified marine, andare almost exclusively are as animals until They 1825. ognized rec not formally to be plants and were long believed were they In fact, orothersubstrates. tothe sea floor attached a motionless and live Sponges are it continues to grow.drifts andsettles to the bottom, where which asalarva andis released velopment inside the parent de by begins budding, casetheembryo sexual,or in which may Reproduction beasexual, the cell membrane. through takesplace andwastes gases respiratory the exchangeof is entirely muscles.cells or Digestion nerve no senseorgans, tract, into cells function without being organized individual in which have a Parazoa support. materials for structural andsecrete foodparticles, filtered anddigest the store transport, currents, the water create thepores, form thesac, of cialized cells coverthe surface Various is pumped and filtered. water spe which through canals of (hence the phylum name "Porifera")with a network zoa belong toSubkingdom all other metazoans who Subkingdom of members andunusual animals. are most primitive They PoriferaPhylum Parazoa the Metazoa. haveclassified study animals) traditionally who ing these body plans, andhow they unfold dur on the basis of plans within these phyla,evolve but they maintain the basic havecontinued to forms New multicellular organisms. large calcareous spongescalcareous embryonic development embryonic radially symmetrical radially (“true animals”). Sponges are basically porous sacs basically animals”). Sponges porous are (“true tissues laid down during the Cambrian Period.It is primarily intracellular and Parazoa organs ( sponges ), silicon dioxide ( (takes place inside the cell) and cellular level of organization of cellular level (“near animals”), distinct from (“near animals”), distinct from sclerosponges . There is no mouth or digestive . There animals, the demosponges ): Sponges are amongthe ): Sponges are , that, sessile zoologists Radiata glass spongesglass life, permanently life,permanently ). ) or acombi (scientists . More ad . More Eumeta body body ), ------, are represented by twophyla, represented are than in sponges andless complex than in bilaterians. Radiata complex these animals is more but body no anus. plan of The a ence of made bycells. is the pres Another advancement the Radiata andectoderm the endoderm from specialized tissues grow or no cells, called the toderm the twotissue layers, are there Radiata, of asfunctional units. In the embryos cells that perform of organization Unlike the sponges,tive. display Radiata a ac allows animals to bemore symmetry bilateral whereas animals,sessile floating or inactive, relatively acteristic of char (asin sponges) are andasymmetry symmetry Radial vanced, and plex life histories,both polyp andmedusa stages including asexually andsexually, reproduce They andmany have com cells,sometimes assembled into sensory net system called a nervous ians also haveaprimitive take placeintracellularly.digestion and absorption Cnidar lular digestion Partial take place. both digestion and circulation coelenteron faster. asthemselves,and much aslarge immobilize prey The stinging cells called equippedwith tentacles, comes tothem,using their sticky theirprey.they cannot pursue food that theyInstead, trap andsenseorgans, nerves muscles, without well-developed but carnivores, in the medusa. Cnidaria are and downward in the polyp isorientedupward the mouth,which rounding polyp the free-swimming occur, Two reefs. body forms the coral form al animals, which tiny cor of flower-like sea anemonesandhuge aggregations a ton and may feet long.100 include have tentacles They jellyfish, the enormous to forms, tiny translucent from allthe animals. range They of als Cnidaria Phylum ): These are some of the most beautiful and important the most beautiful and important someof are ): These , with colonial or , and a middle layer of gelatinous material with few gelatinous material with few , andamiddlelayerof bilaterally symmetrical bilaterally protoneurons hydroid gut cavity functions as a forms. There are about 10,000 known spe 10,000 about are There forms. , where cells are organized into tissues, layers into tissues, layers organized cells are , where by enzymes takes place in the gut, andfinal ( form. Both have whorls of tentacles sur tentacles of Both havewhorls form. medusa sea anemones, jellyfishes nematocysts mesoglea , calleda Cyanea arctica Cyanea (primitive nerve cells) and various cells) and various nerve (primitive or jellyfish form, andthe sessile orjellyfish form, gastrovascular cavity gastrovascular Cnidaria coelenteron . During development, more more . During development, . They can capture and can capture . They animals are animals are , which weighs almost , which endoderm and tissue level of of tissue level sense organs , with a mouth Ctenophora Bilateria extracel and and , where , where nerve nerve cor ec ------. . . 83 Lesson 7: Marine Invertebrates—The lower phyla , ------phy char (with a (with , providing eye-spots , pseudocoe ): Platyhelminthes brain coelomates molecular sequenc morphological flatworms ( , and some have organs for hydrostatic skeleton (without a coelom), (without . This is a second body cavity, located . This cavity, body a second is aremore shedding and subject, on the light coelom relationships. Once again, (with a “false (with and cavity”) acoelomates techniques

is presentis flatworms,in have a and they chemoreceptive organs Their metabolicrate aboutis higher 10 times graspingprey. areRadiatain than and they more active. parasites, Mostly and excretory respiratory, digestive, their are systems primi tive, whilereproductivetheir organs are Theircomplex. quite and flatsize small and their materials, their all supply hosts Traditionally, have been dividedthe Bilateria three into Traditionally, groups: lomates true coelom). Zoologistsused these classifyhelp to acteristics understandand animals their logenetic ing appearsit divisionsthese that are clear-cut.so quite not We use them here,will invertebrateto divide the many groups, (acoelomates and "lower" phyla into somewhatarbitrarily, pseudocoelomates), coveredand "higher" lesson, this in (coelomates), covered phyla in the next. Phylum Platyhelminthes (“flat parasiticworms”) have long been considered the sim bilateralhaving animals plest symmetry and an embryo with three layers. cell Themesoderm to formsplit does not a coe does form lom, but muscle some organs.and Cephalization between the ectodermbetween the and endodermthe developing in em bryo, ofinstead mesoglea Radiata. seen in the gelatinous Themesodermal formcells muscles,circulatory the system, most organsofoutside tract, the digestive verte and (in brates)system. Theskeletal the endodermalform cells the tract,digestive whilethe ectodermal become cells the skin and nervousa third Having system. embryonic oflayer cells greatsecond the also made possible innovationof bilaterian evolution:a wall.body and the cavity gut between the The coelom forms when the mesodermtwo layersinto splits and creates a fluid- cushioned Organsare cavity, the in suspended cavity. filled coe the fluid.the soft-bodied animals, protectedIn and by fluidlomic a also acts as supportThelocomotion. in and aiding development of the coelom wasto the evolution essential of large, ani complex mals. . - - - ) ), - , and , with comb ctenes ) surfaces) Nuda tissues . ” or hydrozoans, cephalization organ systems ventral , a third layer of, a third layer cells combs ): Comb jellies appear jellies Comb ): and was crucialthe coursein anthozoans ) ends, as well as left and as well ends, ) , with tentacles,, with or organs and mesoderm ) and bottom ( and ) surfacesRadiata)also front in but . Food is caught by surface mucus by caught is . Food comb jellies is used for balance. There balance. for used is are only posterior ( dorsal aboral Tentaculata bilateral symmetry and colloblasts ) and rear) ( oral composed ofplates (“ ciliated anterior tissues aretissues organized into other importantTwo advances occurred Bilateria. Thein first wasappearance the of the with the mouth, sense organssense mouth, the with and nerve forward center at the sensory and nervousend. locomotor, As developed, systems more efficientrespiration, circulation,and excretion digestion were required.demonstrate Bilateria levelmost complex the of organization,where are cells organized into right sides. Movement could be directedcould Movement sides. right forward (anteriorly) and became more efficient,to more suitable active lifestyles. Increased the development activity ofinvolved muscles, more organs,specialized sense and a more nervous complex sys same time, a head developedthe tem. At ( The advent of of evolution. animal Thisarrangement to animals allowed top ( only not have the (like ( out tentacles. and Pseudocoe Bilateria—Acoelomates lomates aboral sense organ around arethey ctenophore100 known vora but species, predatorscious numbers and can occur in largeto enough important.be ecologically They are exclusively marine, and are classifiedas cells called cells and carried the stickycilia, or by tentacles by to the mouth acrosswiped being Themouth. the gastrovascular is cavity somewhatmore advancedCnidaria, and a unique in than tentaclesare lackingor are whorlsin not around mouth. the Theyradially-arranged eight areby distinguished rows which areThey lack swimming. for used nematocysts, but tentacles,their whenpresent, areadhesive equipped with cies, almost all marine, whichall almost cies, are classifiedas scyphozoans, cubozoans Phylum Ctenophora stage), but have no polyp to Cnidarian medusae (they similar 84 BIOL F150: Introduction to marine biology these parasites, including long a 13-m nematode that para including these parasites, hosts to are marine organisms of animals,and mostgroups matter. plants and organic parasitize Many roundworms feeding onbacteria anddead acre, per billion individuals eral up to sev they can occur in densities of sediments where mostly in soils and found everywhere, 5 animals. are They every accounting 4out of for perhaps dant animals on Earth, themostwidely distributedsome estimates are andabun and by numbers, seldom seenanimals occur in incredible NematodaPhylum discussedinthenext lesson. are members the superphylum with the studies Nemertea place the proboscis, cavity housing the pulling itinto the mouth.The and retracts, theprey around then wraps toxin. a proboscis secretes The and with a "stylet" that wounds the prey boscis is also armed the prey. that ensnares surface sticky In some types, the pro its cavity, from it inside out and exposing a everted turning a andhave aunique muscular tube called carnivores are They systems. muscular and nervous developed anus, and more globin pigment tem, sometimes containing the respiratory sys circulatory an enclosed including than the flatworms, features advanced havemore They the longest invertebrate. ft) in 100 the case of m (nearly 30 to millimeters just a few in size from range They and sand. orin mucus tubes in the mud and algae among the rocks living waters, but abundant in shallow temperate they are mostly marine, worms, ribbon known 900 species of about Nemertea Phylum thislesson. some smallphyla describedattheendof in the superphylum flatworms found in or on other animals). Many now place the zoologists larians and vertebrates), of flukes, also common parasites most vertebrates), intestines of cluding known, in Platyhelminthes are species of About 20,000 easily exchanged. are nutrients gases, and wastes spiratory ratio,so re theirsurface-to-volume increases tened shape proboscis , acomplete digestive system with both a mouth and (predominantly marine, free-living carnivores, some carnivores, free-living marine, (predominantly cestodes , used to capture prey., used to capture is explosively organ This rhynchocoel Lophotrochozoa (segmented tapeworms parasitizing the parasitizing tapeworms (segmented ( ( roundworms ribbon worms ribbon , is similar to acoelom.Recent Platyzoa Trochozoa trematodes (Fig. other whose 12), Lineus longissimus Lineus ): These mostly small, ): These (Fig.along 12), with ): There are only are ): There , a subgroup of of subgroup a , (leaf-shaped (leaf-shaped turbel hemo ------, dermis longitudinally, sonematodesmove byTheir thrashing. an incompletely lined systems, and organ andreproductive excretory,nervous havedigestive, They food is pulled in and crushed. where riorly, opens into a muscular nearthe mouth which ante concentrated are andabrain both ends.Sense organs at section, tapering in cross slender and circular like") are andsimple body plan.Nematodes(“thread- tively uniform has a decep group successful and diverse enormously This amonganimals. arthropods, -secondonly aheadof)the to(oreven almost unparalleled their diversity, would be true, aswell as their numbers, were this million to 100 If species! 40,000 from range evidence, their diversity, molecular from known, but estimates of are species and 25,000 whales.Between 10,000 sitizes sperm Ecdysozoa. in with and arthropods, nematodes classified been recently these little-known pseudocoelomates have the meiofauna, Phyla Kinorhyncha describedin your textbookare (pp. 128and295). (along with Platyhelminthes) in cluded mayThey haveapseudocoel. in be lomate andtheothers acoe are a lobster’smouth). Gastrotricha around the hairs inhabits which (except Cycliophora, tween sediment particles animals terstitial to as referred and Gnathostomulida, Gastrotricha Phyla Cycliophora, (Fig. andmolecularevidence logical 12). morpho and Ecdysozoa), basedon current Lophotrochozoa (Platyzoa, lar, superphyla different in nowplaced three are although and Nematoda), superficially simi thes,Nemertea animals we havejust described(Platyhelmin worm-like of phyla we will meet in the next lesson.Noticethat the three etc.),whom (insects, crabs, arthropods the hugely diverse Ecdysozoa beenplacedin the superphylum nematodes haverecently molting called is periodically aprocess shedasthe animal grows,

Rotifera (skin) is covered by (skin) is covered atough,elastic . Based on these features, and molecular evidence, and molecular evidence, . Basedonthese features, (“molting animals”). Ecdysozoa also includes (“molting animals”). Ecdysozoaalso includes : These tiny, bizarre-looking creatures are tiny, are : These creatures bizarre-looking meiofauna , which live in the equally live tiny spacesbe , which

pseudocoelom and

(“diminutive animals” ), or (“diminutive Loricifera Platyzoa . The muscles all run allrun muscles The . : Also members of of Also : members cuticle . Meiofauna . Meiofauna pharynx which which epi in ------, , 85 Lesson 7: Marine Invertebrates—The lower phyla - - blasts blasts protoneuron and nerve net, neuronand brain, central nervous system eye-spot hemoglobin proboscis pharynx meiofauna/interstitial animals Cambrian Explosion Cambrian Period, asexual and sexual reproduction, and sexual asexual hermaphro dite and broadcast spawning sessile cells and spicules osculum, ostia, collar hydroid (polyp)medusa and aboral statocyst, nematocyst, cnidocyte, sense organ planula comb rows/ciliary combs, ctenes and collo • • • • • • • • • • • • • • - - glossary terms glossary molting and metamorphosis intracellular and extracellular diges digestion tion filter suspension feeder, deposit feeder, feeder glea gut/digestive and gastrovascular cavity cavity coelenteron, coelom and rhynchocoel hydrostatic skeleton epidermis, gastrodermis and cuticle oral and aboral, ventral and dorsal, anterior and posterior cephalization endoderm,ectoderm, mesoderm and meso acoelomate, pseudocoelomate and coelomate pseudocoelomate acoelomate, cellular level of organizationtissue level and of organization radial symmetry and bilateral symmetry Metazoa, Parazoa and Eumetazoa and invertebrate Radiata and Bilateria • • • • • • • • • • • • • • • • your textbook. your Listed below areListed below terms some key Do a self-check this lesson. for understanding. for If you're unclear about the meaning of ofany these terms, re-read in this course the material and check textbook, guide and in your the glossary at the back of 86 BIOL F150: Introduction to marine biology online at eLearning.uaf.edu and in the course materialsyouonline ateLearning.uaf.edu received. andinthecourse a Include thesubject.) hasnoknowledge of someone who something,"describe" doso-don't just list. (To explaining that you to imagine are andshow your beclear understanding, own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 6. 5. 4. 3. 2. 1. closest relative: flatworms, ribbon worms, insects, you? worms, ribbon flatworms, relative: closest their thefollowing animalsare of Which may animalsonEarth. bethemostabundant Nematodes (roundworms) phyla inyour thatoccurasmeiofauna, description. several dothey Include live? andwhere themeiofauna Who are inthislesson. included phyla theworm-like oneof of Describethebasiccharacteristics creatures. worm-like are Many "lower"invertebrates inanimals. symmetry bilateral of thataccompaniedtheevolution advances Describe several theanimalsincluded. their lifehistoriesandsomeof by describesomehighlights of class, and,for four each classes. Nametheseclasses Phylum Cnidariaisrepresented Define, by thefollowing terms: comparingandcontrasting, k. j. i. h. g. f. e. d. c. b. a. cover sheet cover

, not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , and deposit feeder filter feederanddeposit andstatocyst nematocyst hydroid andmedusa netandbrain nerve andcoelom coelenteron andmesoderm mesoglea symmetry andbilateral radial digestion andextracellular intracellular organization of cellular andtissuelevels andeumetazoa parazoa andinvertebrate vertebrate Written Assignment for 7 Written Lesson Assignment and submit the written assignment to UAF eLearning & Distance Education according to directions found to directions & Distance Education according andsubmit the written assignment to UAF eLearning (Please make your completeandconcise) answers • brief and clear , but, also complete . For example, if asked to asked For. example, if in yourin 87 Lesson 8: Marine Invertebrates—The Higher phyla - . Marine Biol Course Guide

(last page of (last page and submit it to UAF the lesson)

Written Assignment 9th Ed.). found at the publisher's Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Be familiar with the various groups of invertebrates, "higher" and their characteristic features. Be aware of some of the major trends in the evolution of these groups, and their relationships. phylogenetic the information of pages Read this lesson in the following for this (Castro Assignment (below) in C&H & Huber, Complete the Reading ogy Complete the eLearning & Distance Education. 1. 2. 1. 2. 3. 115-149) C&H: Chapter 7 (pp. More information, including interactive flashcards, quizzes, videoclips can be and links, Reading assignment Reading steps for completing the lesson the completing steps for In this lesson, we continue our survey of the invertebrates, the "higher" covering invertebrates - those with a true coelom. this Lesson of Objectives Invertebrates— Higher The phyla Marine 8 88 BIOL F150: Introduction to marine biology used for respiration and to capture suspended food particles. suspended foodparticles. and tocapture used forrespiration calleda structure for isnamed a unique group This first. sider thelophophorates la, the superphylum The Lophotrochozoa—Lophophorates in progress. work a much is still very genetic relationships animal taxonomy andphylo of will our understanding see, ing the major animal phyla.show Also look atFig.a family tree 7.54, the of the characteristics blein your 7.1 textbook toreview Use Ta for bigpicture). the Timescale really in the Appendix (andtheGeologic Guideforthe bigpicture this Course of 12 in Fig. scheme to the animal classification back refer Again, later,anus and themouth forms asecondindentation. from tion. In this initial indenta from with the mouth eventually forming cells inside the ball. Mostanimals are layer of a second forming migrating, cells begin in the ball,into which cells. and multiply, As the cells divide ball of adentappears deuterostomia paths: two evolutionary in the embryo.shape Traditionally, have recognized zoologist way the bodyto take begins us), is named for the particular includes (which Deuterostomia superphylum, third son 7. The andEcdysozoa,in Les Lophotrochozoa of lomate members We12). and pseudocoe mettheacoelomate have already Lophotrochozoa and molecular evidence: on morphological based superphyla, W lophophorates phylogenetic relationships deuterostomes Marine Invertebrates—theMarine higherphyla those with a true those with a true phyla, advanced more covering thegenerally marineinvertebrates, of e continue oursurvey (“second mouth”). The embryo starts asa starts embryo (“second mouth”). The Lophotrochozoa lophophore , Ecdysozoa andthe protostomia , the first indentation the becomes the first , , a crown of ciliated tentacles ciliated tentacles of a crown , coelom and trochozoans includes groups of phy of groups includes Deuterostomia of majorphyla. As we of (“first mouth”) and (“first . They fall into three into three fall They . protostomes . We. will con (Fig. • ------, toprocta by four phyla: represented are simple. They systemsare other organ Most the lophophore. opening near tract plete U-shaped digestive sheathing, and mosthavea protective of form some also have a Lophophorates this feature. All the phyla share in this group taching to the substrate by a stalk-like continuation of thebody by stalk-like a continuation of to the substrate taching sessile, at all are species, 150 mostlyabout marine. Nearly only are tentacles.There with the anus is inside the ringof is arranged digestive tract in that their U-shaped bryozoans lum. As their name ("inside anus") suggests,from they differ but nowhavetheirownphy as Bryozoa, classified have been Entoprocta Phylum the phylum (“outside anus”). cles—hence name, Ectoprocta tenta ring of and the anus just outside, the lophophore’s that the mouth lies inside, such gut is arranged U-shaped Their orcreeping. free-floating, are shapes. A few screw or cork orupright with solid, branching may beencrusting, colonies tissue. Bryozoan of otherbyto each athin strand usually connected zooidsare The the structure. cleaning or oids ( food functions,gathering including different form zooidswithin the colony per typesof extended.Different the which from a compartment forming agelatinousrigid calcium structure, or carbonate secretes called tiny individuals colonies composed of fans, with branching like cnidarian sea called "mossanimals,"many looksomewhat Also the world. throughout abundant in are success. They of long record havea identified, bryozoans fossilspecies,living 5,000 andmanyforms about more Ectoprocta/Bryozoa Phylum ), egg production ( production ), egg zooids ), coelom Entoprocta, Phoronida Entoprocta, . A sheath of tissue encloses each zooid and each tissue encloses A sheath . of divided into two or three compartments, compartments, into two orthree divided ( goblet worms heterozooids

(“ , with the mouth and anus moss animals moss , and ): These tiny animals ): These ), and strengthening ), andstrengthening lophophore Bryozoa Brachiopoda autozo ”): With (or can be com Ec ------. 89 Lesson 8: Marine Invertebrates—The Higher phyla

------se ): Thisfamiliar . The wall body fluid-filled coe . Polychaetes are . Polychaetes hydrostaticpres , and are used for gills , and usually by an additional additional an by usually , and hemoglobin closed circulatory system Polychaeta (one per (one segment), and blood (earthwormskin), and their . These protective layers enclose (polychaeta means “many hairs"). means “many (polychaeta segmented worms hearts ( cuticle , are characteristicof The polychaetes. epidermis Oligochaeta (segmented)organization, and a distinct with (leeches) and bristleworms parapodia , make annelids powerful swimmers, make annelids and burrowers. . The tract digestive and organare systems relatively , sense organstactile and are fewoligochaetes Pairedmarine leeches. and arm-like of extensions muscular wallbody the on each segment, called parapodiasupports, have skeletal called bristles tae locomotion. The setae give groupthe their common name, plexity in animal evolution.animal in The plexity plan of body has a annelids metameric dividedmany into head, length and a body along its complex compartments. Eachsegment a contains lom well-developed, includinga vessels, blood with respiratory pigment, usually the coveredis by the outer covering, two musclelongi and inner layers, an outer circular layer each in segment. Thislayer tudinal arrangement of mus cles, and a fluid-filledcoelom to exert sure three major classesof have been annnelids Traditionally, recognized: Hirudinia the largest group and almost all marine, whereas there probably have a common ancestry. Until recently, annelids recently, Until probablyancestry. a common have were considered more closely relatedarthropods, to partly formsadult because their are both segmented.anne But trochophorehave lids larvae, and arthropods don't. Using recapitulationthe morphologicaland other principle, and evidence, sequencing molecular most zoologistsclasnow as trochozoans,and molluscs sify annelids Superphylum in Lophotrochozoa,place arthropods and Superphylum in evidence Similar places the ribbonEcdysozoa. worms (Ne mertea), fromprevious the trochozoanthe in lesson, group. Phylum Annelida includingearthworms phylum, and leeches, importantis for severalperhaps with 20,000 known reasons. large A group, are annelids species, important ecologically members of They arecommunities. sediment and marine soil one also of groups,studied most ofthe models as developing com , ------) is Re are forms phylog Lingula . They prefer ontogeny ): Only about20 Only ): , like the unrelatedthe , like inarticulate ): The): approximately pedicle , plus several, plus smaller valves Trochophore larvaeTrochophore Mollusca (ribbon worms), whom we met lamp shells , or “ontogeny recapitulates, or “ontogeny phylog ( horseshoe worms and ( brachiopodsarticulation have a hinge-like Nemertea Annelida , the other lophotrochozoan group, includeslophotrochozoanother , the two group, Articulate ) of an organism,ontogenies similar and species with eny shaped gut. Trochozoans are shaped illustration gut. Trochozoans a good of the Principle capitulation other words, In embryonicdevelopment ( eny.” a condensed versionof evolutionary the history ( in the previousin trochozoansadults, lesson. As appear may earthworms unrelatedcompletely (e.g. but and octopuses) larvaetheir are nearly identical. bands two cones placed base to base), with (like biconical ofaround cilia the middle, a topknot of flagellae and a U- Lophotrochozoa—Trochozoans Lophotrochozoa—Trochozoans Trochozoa large phyla, including phyla, some formsresemble a type of the hence lamp, oil ancient Most arecommon name, “lamp shells.” attached to a sub strateof means by or stalk, a fleshy cold water the deepthe poles or near in sea. environments, known. Brachiopodsknown. areto aboutup small, and are3 in, pro or halves, two with a shell tected by clams. valves,the between (joint) whereas in valvesthe The muscles. areof shells by held together only Phylum Brachiopoda 350 living species of brachiopods appear vestiges to be the ofmore with a once widespread marine group, and abundant One genus, identified. species 30,000than fossil includes unchanged)animals evolutionarily (most oldest the to aggregate, the tubes twisting around each other. Theto aggregate, around the tubes twisting eachother. name "horseshoeworm" comes fromhorseshoe coiling the shape whichin the lophophore tentacles are often arranged. except polar areas, and may have been morebeen have areas,polar except may and prevalent the in past. Phoronidsare worm-like,and slender from a fewmil limeters Theyburrow to about long. a foot sediment the into rocks, into holes or dissolve and even shells cement piers, secrete.a tube they the hole with line and then They tend wall, topped by a cup-shaped ("goblet") body. Some species Some body. a cup-shapedwall, ("goblet") topped by areindividualsmultiple with colonial, on branching stalks. Phylum Phoronida living occur throughout arespecies they world the but known, 90 BIOL F150: Introduction to marine biology are sedentary, living in burrows and crevices, or the shells of orthe shells of sedentary, and crevices, are in burrows living systems, andmost simple organ have relatively Echiurans extendedused in tobesensation greatly and feeding.be and can (hence forked, their common name)or shaped mayspoon- proboscis be means“spine-tails.”The which thebody,end of the posterior the phylumgiving its name, proboscis plump, body, sausage-shaped unsegmented with a grooved havea They all marine organisms. known 150 species) are Echiura Phylum in progress! chaetes. Taxonomy ascientific work is certainly poly may from andleeches, oligochaetes derived be as annelids, andthat classified the other annelid classes, andSipuncula (described below) should also be Echiura suggests that the phyla and molecular research logical lids,in the family themasanne their own phyla, studies confirm but recent oftenplacedin are Pogonophorans and vestimentiferans example of gut - agreat embryonic called a ized organ synthetic bacteria by butto beprovided the tentacles, most appear through absorbed in the adult.Somenutrients are tract animals withdigestive nomouth or the only free-living are spectacular example (C&H Fig. p. 16.28, worms Beard 380). a vents,are hydrothermal around living giant tubeworms of floor. onthesea carcasses atwhale even vents, and atmethane seeps hydrothermal ments, around the dominant to be appear which thesetubeworms, about lot tolearn collected specimens. We previously still have a of off torn opisthosoma (called the their bodypart is the posterior of mented part canbeaslong as 7ftandthe onlyThey seg recovered. intact specimens were the first as annelids until when 1964, notrecognized Pogonophorans were sea. in the deep live they Completely 1900, unknown before worms). ("garment" worms ( ing worms andbeautiful tube-dwell the bizzare Two are unique groups lifestyles. marine habitats, with many different many different in living Polychaetes abundant extremely anddiverse, are ” - for their long tentacles) and for” - their longtentacles) extending beyond themouth,andsmall hooks at tubeworms macrofauna ), which is buried in the mud andhadbeen ), which ( Siboglinidae spoon worms trophosome living inside the worm in a special inside the worm living ), the of the deep-sea, living in sedi living the deep-sea, of pogonophorans . In fact, current morpho current In fact, . , which develops from the from develops , which ): This small group (about (about small group ): This obligate symbiosis vestimentiferans Riftia , a genusa , (“ chemo beard beard ------.

Phylum Mollusca Phylum apeanut. the animal resembles muscles, by retractor into the trunk is retracted When the introvert trunk. the body into an anterior part, ing on detritus. is divided The or otheranimals’ shells, andfeed crevices in burrows, living worms simple, unsegmented small, relatively of known (320 species,this all is a smallmarine) group Sipuncula Phylum polychaetes and fish. as small crabs, such sal organisms, “ plankton. Commonly calledthe amucus net fortrapping crete tose anduses theproboscis burrow in a U-shaped lives California, thePacific coastof of in mudflats other animals,on detritus.and feeding One speciescommon the mouth. They havean the mouth. They a digestive system is complex, with used in locomotion. The on the developed is by which muscular body wall coelom is reduced athick, and their has specializedsense organs, head developed in the a by means of draw carbonatecalcium tissue called the layer of by meaning “soft”) covered a aword from lusc” is derived havean chozoans.They other tro distinguish molluscs from Many specificfeatures as food sources. their importance becauseof conserved byand they people, havebeenextensively studied and collected andused shells have long been admired, whose oftenconspicuous creatures, phylum. are They vertebrate any otherin than of this group of completepicture more in Wecommon. luscs still have basicfeatures may havea since. However, paths ever evolutionary individual all mol the of beginning the before established even molluscs were of major groups nematodes).The with the possible exception of (again, in marine environments diversity,and is the mostdiverse nematodes)in worldwide (andperhaps only toarthropods as manyknown species, as200,000 this phylum is second innkeeper worm radula introvert mantle cavity (a hard, toothed instrument for rasping food) in for rasping toothedinstrument (ahard, , with a mouth and tentacles, and a plump and a with mouth , a and tentacles, Cambrian Period ventral ,” its burrow may be shared by mayshared commen be ,” its burrow (snails, octopuses, clams, etc.):With retractor muscle retractor shell into which theanimal can with shell into which ( , usually through usually, through peanut worms unsegmented open circulatory system open circulatory (belly) sideasamuscular mantle . The mantle secretes a mantle secretes . The , and they have pursued and they, havepursued . Respiration occurs occurs . Respiration

soft body gills ): Like Echiura, ): LikeEchiura, Urechis caupo Urechis . Their well- Their . (“mol with a foot ------,

91 Lesson 8: Marine Invertebrates—The Higher phyla

. , ------in and seg cara where on the on gills smooth is hard ) or other nauplius open cir trilobites thorax mandibles , or , jointed ap and fused in and , branchinginto hemocoel head , includesspiders maxillipeds tracheae pereopods exoskeleton or Chelicerata maxillae , has a huge varietya huge , has of marine calcium carbonate arepredominant the repmarine , like ours., like Thereduced,coelom is , whichsurface, on the skates the is . Respiration is by . Respirationby is biramous appendages . Fertilizationinternalis most groups. in and were and previouslyto be related thought Crustacea (arthropodaliving In foot”). “jointed means Halobates (one more (one ofa set insects), than regions.of Movement appendages the con is (crustacean insects, Unlike ones”). means “shelled striated muscles striated and two or threeand two of sets head, several pairs of walkinglegs ( types of specialized appendages on the thorax, and usually reduced appendages the abdomen. on These appendages areto perform specialized a variety of including functions, capturefood chewing, and manipulation, walking, swimming, breathing,egg copulation, and sensory brooding, reception. Crustaceanshave a special larval form called the trolled by a complex muscular system, includingmuscular a complex trolledby and and most of open an is cavity body the are the tissues an in blood, bathed directly by culatory system forms.marine The nervousanne an to similar is system a more with brain. but complex lid’s, and Most have single eyes compound Subphylum Crustacea resentatives of arthropods, the 68,000 approximately with identified, which be a fractionspecies only may of true their In most crustaceans, the diversity. ofaddition the ened by the head and thorax region, formingcalled a a shield pace crustaceanshave and outer arms.inner Theirappendages includeoftwo sets antennae ter strider 344). 15.17, p. Fig. ocean (C&H open the in insect only The second, forms, including crabsfamiliarthe lobsters, and shrimp, barnacles. third A group, and scorpionsforms, and some marine horseshoe the crabs and sea spiders.fourth A the group, arthropodother many possibly and are lineages, extinct. arthropodsexoskeleton, to their addition In have a mented body to the segmented worms,primitive In forms, annelids. the each segment was a pair of equipped with pendages arthropods, of many appendages these are modi highly fied or lost, and segmentsare fused into abdomen

. ------and nerve hemo ). These exoskel Uniramia Kinorhyn chitin oflarval the hemoglobin (snails, limpets, limpets, (snails, and molting (clams, oysters and , with about, with 800,000 , forming an = , the roundworms, plus , and an extensive extensive , and an . Thelargest, , a nervous usually system, Loricifera (insects, crustaceans, (insects, spi (squids, cuttlefishes, octo cuttlefishes, (squids, cuticle insects Gastropoda Bivalvia ecdysis is a taxonomic leveltaxonomic a is created metamorphosis ( Nematoda subphyla and the respiratoryand the pigment kidneys (insects, crustaceans, (insects, spiders, etc.): Arthropoda Cephalopoda of molluscs, exoskeleton . Variations of . Variations features these arethreethe seen in (giving the blood a greenish (givingthe blood or color) classes composed oforganic material (usually nerve ganglia, nerve cords (Fig. 12). We covered the last threeLesson 7. in 12). We phyla (Fig. centipedes, and has fewcentipedes, marine representatives. The wa the plant kingdom, insects are kingdom, insects the plant relative late-comers which diversified on land. Theexplosively livingarthropods may be dividedseveral into (unbranched appendages)and includes millipedes insects, More species have been identified in this phylum than in any Moreany in than phylum this species have been in identified The vastaremajority other. more and probably many species, known undiscovered (es rangetimates oftens the into environ marine In millions!). are insects rare.angiosperms the Like however, ments, of ders and relatives) and several including phyla, minor cha Phylum Arthropoda whereasto dramatianimal the for possible makes it molting this form. alter its have exploited cally particular, in Insects, opportunity undergoing by formsvery into different adult forms. include Ecdysozoa major phyla, two other substances), whichother substances), animal as the shed periodically is Thisgrows. the size of to limit tends also but the animal, mineralwith animals In opens other developmentaloptions. vertebrates,and molluscs like skeletons, growthoccurs as moreonly mineralform,animal’s the added, is limiting have common ancestry, based on morphological and mo have common ancestry, evidence.lecular sequencing that animals It comprises the shed their have a three-layeredanimals eton Ecdysozoa Superphylum Ecdysozoa recentlyquite to group together severalto thought phyla major abalones and nudibranchs), and mussels) whichNautilus), and puses are textbook. your described in 2-chambered heart cyanin There are one or two with network 92 BIOL F150: Introduction to marine biology Small, torpedo-shaped and nearly transparent, they dart they dart transparent, and nearly Small, torpedo-shaped should be classified. creatures how these strange ally clear Chaetognatha Phylum and worms), mata phyla include into the Grouped asecondindentation. from mouth forms the embryonic which animal phyla are remaining embryo. The the of in-pocketing thefirst from themouth develops which are so far wehavedescribed groups The Deuterostomia this group. of marine representatives the only remaining now extinct, with are totheCambrian datingback mites.Many marinechelicerates of posed digestive enzymesinto their prey. is mostly com group This unable toingest solidfoodand insteaddrinkblood inject or are Mostchelicerates dibles used by mostotherarthropods. man the chewing food,insteadof usedtograsp are which named for called their pointed appendages, are They andCrustacea. in addition to Uniramia thropods, Chelicerata Subphylum class, A third tothesubstrate. head-first attached sessile lives they live bivalves, that resemble in this class.Shelled forms will see later in this course. levels, aswe and animals at higher trophic the producers vital link a between on phytoplanktonfeeding and forming thesea, of thegrazers are They abundant animals on Earth. plankton socommonamongthe are which tiny crustaceans fishes. A second class, and for food birds somewhales, source and the exclusive abundant extremely as zooplankton in polar seas are which e.g., uitous and important, Smaller, and crabs. ubiq are inconspicuous malacostracans cious) the largest, subphylumsix into about is divided This (sea stars, sea urchins, etc.), seaurchins, (sea stars, Branchiopoda Order Decapoda Order arachnids that they also are contenders for the title of most forthe title of contenders thatthey alsoare Malacostraca Chaetognatha Chordata Superphylum Deuterostomia Superphylum , the terrestrial spiders, scorpions and scorpions spiders, , the terrestrial horseshoe crabs horseshoe , includes fairy shrimp and water fleas. shrimp and water fairy , includes Maxillopoda (vertebrates and their relatives). andtheir relatives). (vertebrates ( ("ten-footed") - lobsters, shrimp ("ten-footed") -lobsters, is the third group of living ar living of group is the third amphipods, isopods , includes the familiar (and deli the familiar , includes arrow worms arrow (arrow worms), worms), (arrow Barnacles Hemichordata , includes , includes deuterostomes and also are included included also are protostomes classes ): Itis not re sea spiders Echinoder chelicerae copepods and . By far . By far (acorn (acorn , these zoo krill , in , in ------, , ,

mary bilateral symmetry bilateral mary from the only animals to make this switch are They development. embryonic pathof low the deuterostome fol which symmetrical larvae, havebilaterally echinoderms symmetrical as adults, etc.): Although radially cucumbers, Echinodermata Phylum shallow seas. especially in warm numbers, cur in staggering onlyspecies, 100 but about are they can oc There wall. thebody through directly andwastes and exchangegases cilia, the bodymove cavity by materials through means of and have norespiratory, organs, excretory, or circulatory by not in use. They covered when ahood andare prey ture shut to cap snap thewhich head, spines extending from eyes system,and andnervous muscles developed swiftly,about have side and tail They like arrows. a head, and theya head, havean they are Radiata, primitive more spines into various modified are which microstructure, tiny skeleton of is made and the endo is uniquely into complex divided partitions, carbonate endoskeleton andcoelomdevelop, andin their the mesoderm in how some similarity bear to chordates, sensation. They tube feet system, the petalson a sand dollar.or the five unique hydraulic Their astarfish of arms can be seenin the five systems, which the skeleton and mostorgan of organization five-fold have a are Echinoderms size. comparable of than any otherorganism urchins heart and seastars more are forthe BeringSea, example, there of In parts and can be extremely abundant in some areas. today still widespread than in the past, they are noderms echi fewer are Although there million years. past 550–600 Period during the many andhave unique developed features the Cambrian of well established near the beginning were fossil all marine. They species described, thousands of of species,cessful phylum,living andtens 7,000 with about symmetry. bilateral is an oldandsuc This to atertiary metry . Voracious marine predators, they haveteethand Voracious. marine predators, in the adult. Some echinoderms even switch again, again, switch even in the adult.Someechinoderms (echinodermata means“spiny-skinned”). Like the (echinodermata which are used for movement, respiration and used for movement, respiration are which water-vascular system water-vascular pentaradial ossicles (sea stars, sea urchins, sea sea urchins, (seastars, oral . But the echinoderm coelom . But the echinoderm to (or surface where the mouth where surface secondary radial sym radial secondary with a unique sponge-like unsegmented pentamerous , hasextensions called compound plates fins and lack andlack calcium ) - they , well- and pri ------93 Lesson 8: Marine Invertebrates—The Higher phyla , . ------ sea ” or ” also , the tunic and Salps house Pyrosoma , retain the ): Thissmall into the sim the into Urochordata ): Appearanc vertebrates where water en . In lancelets Larvaceans . At some point in their in some point . At wherewill (and exits it tunicates ( ( salps metamorphosis arefilter-feeders, sessile simple Vertebrata notochord,nerve cord and incurrent siphon and , a column of, a column articulatingbones or cartilage excurrent siphon ). They). are ofsubjects the three next the lessons. sea squirts , sometimes seen in the warm, sometimes seen in waters off New larvaceans , . The name Urochordata (urocomes from = tail) the backbone , sometimes more filter size, for ten times their body than vertebrae to 60 feet long! Tunicates are widespread and often abun to 60 feet Tunicates long! can they muchinsights dant, and have been the for studied provideancestrythe into ofchordates, higher ourselves. like Subphylum Cephalochordata impor about also is two dozen species, only with group, clues for to understanding primarily tant developmentthe and evolutionof higher chordates. eel-like animals, Small are for tail muscular a strong, with tadpole-like animals swimmingand a clear slits factthe notochord that present is the tail. The in only lar vaesea bed, on the attaching soon settle head-first and undergoingdramatictheir form.ple adult The notochordare and tail absorbed, and the sieve-likebecomein the openings slits the gill tunic. transparent called tunicates, Tiny, pelagiclarval form. Theysecrete “ a gelatinous net particles.out food ing becomes When the filter clogged, they Newarenets secreted rapidlyescapecan througha “door.” every few hoursand frequently, some species. in arebarrel-shaped with tunicates, planktonic bodies within transparent Some are tunics. including colonial, spinosum sock-shaped luminescent, brilliantly in Zealand up colonies Three share subphyla of all traits: these Cephalochordata embryonicnotochord surrounded becomes and replaced a by ( Subphylum Urochordata es are deceivingthe case of in urochordates, or tuni They includecates. about 3,000 marine species of squirts are they encloseda sac called a in history, life adults, As tu Enclosedslimy their in resemblinganimals sponges. have an they nics, ters, and an squirt out if are they squeezed!). Their larvae, however, ------gill ver , and , which and of chordates, - each distinc and relatives): zooplankton ofchordates. The stomochord ) - a series of) open - a bundle of - a bundle nerve fibers, Crinoidea nerve cord surface. Most organ systems acorn worms notochord lifestyle as adults. This as adults. lifestyle combi and arefrom distinguished ( Asteroidea,Ophiuroidea, Ech tunicates, lancelets tunicates, aboral - an extension of - an extension wallbody the mus ( benthic pharyngeal slits - a flexible cartilaginous - a flexible rod ly Chordates (or ): notochord post-anal tail (some fossil echinoderms fossil (some ofsigns show also slits), gill dorsal, hollow nerve cord a underneath,ing supporting,and nerve the cord. a culature, nerve cord and notochord, behind the anus. a often enlargedforward at the end to formbrain, the and runningalong the back pairs with of nerves branchingoff at intervals,the brain connecting to the muscles and organs. gill slits ofinside the connecting ings throat the ex to the filter feeding. and for often used as gills terior, presentdevelopment: their during at some time DNA studies, also show a closeshow also affinitystudies, echinoderms.the DNA to Phylum Chordata tebrates features, four each ofby other animals which is slits a structure reminiscentof the and a rudimentary structure called a somewhatresembles the larvae of some hemichordates, and the resultsof some our own group, the chordates, and other animal phyla suchchordates, the phyla other animal and group, our own as echinoderms. Hemichordates (“half chordates”)share ofall, not some, but chordate the features. Theircoelom deuterostomes, in develops and is di only seen a in way threevided into regions,vertebrates. as in They have Phylum Hemichordata Thisof phylum small worm-likecreatures infrequently is livingseen mud, or among rocksthe in tubes in sea and ofis weed. It between link interestas a possible primarily echinoderms one of Theythe most distinctive phyla. animal includefive classes- the Holothuroideainoidea, tive own right, whichits in are textbook. your described in lies, and an opposite lies, the areexcretory and simple, organs are absent.Typically, larvae swimabout as part of pelagic the a into settle then ofnation primitive, advancedfeatures and unique makes 94 BIOL F150: Introduction to marine biology a backbone. The best known lancelets are a group called agroup best known lancelets are The a backbone. and they lack developed, poorly are and sense organs is tiny their brain into the head. But, unlike vertebrates, that the the fact = head) comes from (cephalo nameCephalochordata life. The throughout oped slits with the head poking up, their through filterfeeding buried in shallow, sandythey environments usually lie half . The other chordate features also are well devel also are features other chordate . The notochord extends gill - of vertebrates are the subjects of the next three lessons. the next three the subjects of are vertebrates of is still unclear. in general) classes chordates different The (and of vertebrates of and the ancestry oversimplification, however,an tobe information, appears this once again legendandsong!).With more (becoming the subject of evolved higher chordates which from form the archetypal amphioxus (“pointed at both ends”), once thought to be 95 Lesson 8: Marine Invertebrates—The Higher phyla - water-vascular system and tube feet tunic protochordate gill/pharyngeal slits, nerve cord, notochord and post-anal tail backbone (vertebrae) siphon proboscis compound eyes biramous opisthosoma and trophosome foot, radula, threadsmantle, pen, byssal respiratory hemoglobin pigment: and hemo cyanin gills and tracheae • • • • • • • • • • • • • - - - glossary terms glossary parapodia and pereopods setae open and closed circulatory system carapace and os exoskeleton, endoskeleton, sicles epidermis and cuticle chitin and calcium carbonate metameric/segmented and pentaradial/pen tamerous ecdysis/molting and metamorphosis symbiosis obligate lophophore trochophore larva and tadpole larva and phylogeny ontogeny principle recapitulation protostomeand deuterostome Lophotrochozoa, Ecdysozoa Deuterosto and mia • • • • • • • • • • • • • • • your textbook. your Listed below areListed below terms some key Do a self-check this lesson. for understanding. for If you're unclear about the meaning of ofany these terms, re-read in this course the material and check textbook, guide and in your the glossary at the back of 96 BIOL F150: Introduction to marine biology online at eLearning.uaf.edu and in the course materialsyouonline ateLearning.uaf.edu received. andinthecourse a Include thesubject.) hasnoknowledge of someone who something,"describe" doso-don't just list. (To explaining that you to imagine are andshow your beclear understanding, own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 6. 5. 4. 3. 2. 1. Briefly describe the four distinguishing features of chordates. of Briefly describethefour distinguishingfeatures separately) group (discuss each andtunicates. echinoderms trochozoans, thefollowing groups: of each thephylogeny of we canbetterunderstand principle, principle?Explainhow, therecapitulation andapplying What istherecapitulation by studying theirlarvae animals belongs. these similarlifestyles?Explain your answer, of each identifyingthephylum becauseof including towhich structures oristhisacaseof related, closely are organisms by plates. protected Doesthismeanthatthese calcareous alsoare andbarnacles carbonate shellwithtwovalves, Lingula other? they toeach belongto, are related andhowclosely phylum organism Which doeseach ference. for feeding, ways. andconeshellsuseaproboscis but Explainthisdif they useitin different worms Both innkeeper subphylum/class.belonging toeach inthatphylum, Namethemajorsubphyla included andsomeorganisms orclasses scribe itsprincipalcharacteristics. phyla, de andbriefly important themore inthislesson.Chooseoneof covered animalsare of groups Many diverse Define, by thefollowing terms: comparingandcontrasting, j. i. h. g. f. e. d. c. b. a. cover sheet cover

, not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , mantle andtunic worms andhorseshoe crabs horseshoe molting andmetamorphosis andcuticle ossicles andmetameric pentamerous tube feetandmollusc'sfoot andpereopods parapodia worms andacorn peanut worms andEcdysozoa Lophotrochozoa anddeuterostome protostome , barnacles and clams look remarkably alike at first glance,since alikeatfirst lookremarkably andclams , barnacles Written Assignment for 8 Written Lesson Assignment and submit the written assignment to UAF eLearning & Distance Education according to directions found to directions & Distance Education according andsubmit the written assignment to UAF eLearning (Please make your completeandconcise) answers • convergent evolution convergent brief and clear , but, also Lingula , in which unrelated species evolve similar speciesevolve unrelated , inwhich complete and clams haveacalcium andclams . For example, if asked to asked For. example, if in yourin - - 97 Lesson 9: Marine Fishes - - . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). found at the publisher's Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Be familiar with the major groups ofaspects of some fishes, their evolutionary his and the features distinguishing them as vertebrates.tory, Understand adapted how fishes are to their environ and physiologically anatomically ment. the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education.. 1. 2. 1. 2. 3. 151–175) C&H: Chapter 8 (pp. More information, including interactive flashcards, quizzes, videoclips can be and links, Reading assignment Reading steps for completing the lesson the completing steps for Fishes are the oldest and simplest vertebrates, and gave to all the other vertebrates. rise They remain the most numerous and diverse vertebrate and are animals, the most importanteconomically marine organisms.. this Lesson of Objectives marine fishes 9 98 BIOL F150: Introduction to marine biology Kingdom Metazoa Kingdom EukaryaDomain Phylum Chordata Subphylum Vertebrata Gnathostomata (jawed vertebrates) Agnatha (jawlessvertebrates) bowfins gars Neopterygii (bony fishes) fishes) (cartilaginous Lampreys Sarcopterygii (lobe-finnedfishesandtetrapods) Sarcopterygii fishes) (ray-finned Actinopterygii skates, rays , Elasmobranchii: chimaeras/ratfishes Holocephali: Fig. Classification 13fish Chondrostei: sturgeons, paddlefishes, birchirs Chondrostei: tetrapods lungfishes coelacanths teleosts: mostfishes, perch e.g., ,, 99 Lesson 9: Marine Fishes

. ------cal Silurian to evolve. mammals fishes, pos fishes, , the “four- , the exoskeleton bones, teeth bones, jaws and jawless and an , a well-defined head, tetrapods fins and and other endoskeletal struc and other endoskeletal , much previouslyany older than fishes skull endoskeleton in their bodies to formtheir bodies in , whichprovides support and protection, , and 50 million years, and 50 million more for amphibians, reptiles,birds scales for some fishes to developfor the abilityto accumulate cium phosphate and wereFishes relatively raretime, whilethis during inverte brates flourished.Then a profusion of both a bony sessing of appeared plates or scales, bony the during arevertebrates, the oldest and simplest and gave rise to vertebrates.other all 8.1 13 here,Fig. and to Fig. Refer ofoutline an for textbook, your in classification, fish and end oftableat the the use Chapter 8 to reviewcharthe ofacteristics various the groups oflook at the Also fishes. Geologicthe Appendix Timescale, in ofCourse this Guide. First Fishes Thewere fossils fish oldest known discovered just 1999, in rockssome 530 million-year-old in ofsite at the China, in sea bed. Thisan ancient oftime at the places them the Cambrian Explosion vertebrateknown lacked Paper-clipsized, they a fossils. had other features but skeleton, teeth and jaws, set bony apartthem ting from lower chordates, includingsophis and musculature,ticated gills and a paired eyes, tures made of cartilage.years30 million took another It endoskeleton morefor and allows efficientvertebrates.in locomotion Vertebrates include footed” Vertebrates far are the most diverse by group of chor ofthe subject and fishes, dates, lesson, are far this by the most numerous and diverse of the vertebrates. They , , ------, a • . Thisse ofap the vertebrae , also called marine fishes : the tunicates : the also evolvedalso to , called skull dorsal nerve cord . The feature key is that Subphylum Vertebrata vertebral joints , a cartilage , we learned, includessub 3 vertebral column protochordates or , whichencloses and protects the . Additional bony or cartilaginousbony . Additional appendicular skeleton appendicular gill slits bone post-anal tail post-anal spine

or and a Phylum Chordata axial skeleton

backbone the most diversemost the organisms, multicellular in and that vertebrates most offor account with that diversity, In the previousthe In learned we lessons, areanimals that protect the nerve cord wherethe head into to it extends make ribs) form brain. the The (and, later, and spine skull up the the make up elements form they the pendages (paired Together, or fins limbs). elements made made elements of whichare separated mobile by ries of vertebrae forms the the nerve cord.or cartilaginous bony A Thenotochord a rod is derived from mesoderm lying cells, nervethe between cordstiffis It gut. the and for enough more endo rigid the than but muscles flexible to attach to, of exoskeletons and skeletons non-chordates. ver adult In tebrates,notochord the replaced is of a series by hollow vertebrates 4 features display set chordates that apart from other animals: notochord vertebrates,in unique them apart setting fromother ani all a result is mals, of further the evolution of notochord. the We alreadyaboutsome know things vertebrates.We know We aretissues their that organized highly organs into and or are they gan bilaterallysystems, have a true symmetrical, that also know coelom, and develop We as deuterostomes. phyla, 2 of 2 phyla, which are The lancelets. and third group, other 3% ofthe constitutes the is and species animal ofsubject the remainingPartin lessons 2 ofcourse. this more 97% than of surveyed We some ofspecies. animal among the 35-40 currentlythe major phyla, animal de scribed. I 100 BIOL F150: Introduction to marine biology tilaginous fishes tilaginous totheir skeletal material, according into two groups, jawed fishes both the including (“jaw-mouth”) forall other vertebrates, by the egories, into two major cat divided Traditionally, are vertebrates The Family Fish Tree all marine organisms. of the most economically important and all vertebrates, and abundant of the most diverse far fishes,the ray-finned of branch At the sametime,asingle declined. fishes havealsogreatly and by the two small groups, now represented are once-abundant jawless fishes The million 50 years. about lasting only the Devonian, extinctbecame at theendof placoderms highlymya. successful 370 and diverse The rise tothe first and gave the Devonian copterygii ray-finned ( fish group, the other modern of resentatives skeleton. Rep cartilaginous to a laterreverted which derms bony placo from to havedescended believed are nian. They fishes the group of Ancestors fishes. freshwater the first someof included placoderms edges. Devonian self-sharpening sometimes with razor-like, teeth,they hadbony platesonthejaws, Instead of feet. 10 than to more somegrowing forms, ing many heavily armored during this includ period, prominent (“plate-skinned”) were rapidly. and they diversified predators, tive of evolution The mya),(415–375 sometimes called“ during the jawed forms expansion of and eventuallyextinct, became with the declined derms feeding on detritus and small slow-moving prey. Ostraco slow, probably foot long and were animals, bottom-dwelling tracoderms Period bony fishes hagfishes ), including sharks, also flourished during the Devo also flourished sharks, ), including (450–415 mya). These armored forms, called forms, armored mya). (450–415 These hagfishes Agnatha ) forms. Lobe-finnedfishes dominated during ) forms. ( (“shell-skinned”), were generally lessthan a generally (“shell-skinned”), were ) also appeared at this time, including both atthis time, including ) also appeared Actinopterygii and the , and cartilaginous fishes and lobe-finned and cartilaginous , jaws ("without jaws"), represented today only ("without jaws"), represented and ( Chondrichthyes allowed these fishes to become ac lampreys Chondrichthyes tetrapods ) and teleosts , and , The Age Of Fishes AgeThe Of . Fishes with jaws fall . Fishes with jaws fall lobe-finned ) and Devonian PeriodDevonian tetrapods , have become byhave become , Gnathostomata ( cartilaginous cartilaginous Osteichthyes bony fishes Placoderms lampreys ( , about about , Sar car os .” ------

Agnatha Agnatha—Jawless Fishes made by evolution these groups. in vertebrate the advances today. alive brates) highlights of some following survey The fishes (andverte marineandfreshwater majorityof vast teleosts.plus one huge group—the fishes, mostly primitive, freshwater of small groups several forall other bony, jawedfishes. and thetetrapods; of relatives the primitive copterygii rays fishes, and two subgroups, ( sensory organs: paired eyes lost secondarily), (probably paired an organs: sensory typesof and three brain a five-part supports which shelf in that theypartial havea chordates than lower advanced young. more miniature bear are They which eggs lay leathery stage, hagfishes large have no larval Unusual in that they theirownsecretions. their nostrils of must sometimes “sneeze” just to clear these lovely creatures months without eating.up toseven Also called and can go haveaslow metabolism abite.They to tearoff flesh a hunk of selves into knots, sliding the knot against with tooth-like is equipped the mouth, which surround Sensing tentacles theinside out. right in and eatingfrom sometimes diving or dying and dead fish, rine fishes dine on polychaete worms Hagfishes altogether. Subphylum Vertebrata oftenleft out of fishes are tojawedfishes than tohagfishes,and hag related closely now thought to bemore these similarities, are lampreys byDespite parasites. scavenging, are and many lampreys their circular, exist muscular mouth. Hagfishes mostlyof sometimes called are slime. They with smooth skin is covered skeleton. Their a cartilaginous andthey fins, have scales and astomach, paired also lack species between them.Besides 50 being jawless,about they animals including Osteichthyes . Bony fishes also are divided into two groups: into two groups: divided Bony. fishes alsoare are the most primitive living fishes, living eel-shaped the most primitive are : Pinkish : and almost blind, ma theseexclusively (“flesh fins”) or (“flesh Elasmobranchii coelacanth rasps ). Chondrichthyes are further divided into divided further ). Chondrichthyes are Holocephali hagfishes for tearing flesh. They cantiethem They fortearing flesh. and lobe-finned fishes for the lungfishes for the and Actinopterygii Ray-finned fishes Ray-finned cyclostomes Teleosts lampreys skull sharks, skates sharks, chimaeras , the closest living living , the closest , a cartilaginous a cartilaginous , makeup the (“ray fins”), (“ray slime eels slime , with only , including , including , because or rat or include include Sar and ------, Lesson 9: Marine Fishes 101

------). cra cte , plus pelvic sharks, or fin rays arewell ( . Cartilagi lungfishes swim blad swim . The Holocephali cycloid spiracle (side fins towardfins (side the ray-finned fishes ( ). There). aregroups two (bones in the inner ear). the inner (bones in lobe-finned fishes lobe-finned (shoulder bones), (shoulder Elasmobranchii ( ventral mouth group, with more with 20,000 than group, which are lighter than water. whichare water. than lighter system for controlling buoyancy. Their controlling for buoyancy. , which includevastthe majority ) and pectoral fins homocercal , largehead ( the bones covering teleost , and a Actinopterygii formold with mouth, the in continuously . Ray-finned fishes include fishes . Ray-finned several small ), which are textbook. described your in is positioned at the front of the head, and oil bodies tail ratfishes Sarcopterygii pectoral girdle ) ) and bony fishes bony in males, whichmales, in in for a penis like function rays teeth swim bladder swim or , a bony plate covering and protecting plate covering gills, , a bony the ), a and rooted teeth scales, made of scales, bone and coveredand mu skin with claspers coelacanths (side fins towardfins (side rear the ofare body) the modified , and lift is provided by their streamlined their provided, is and lift shape, by pectoral tetrapods heterocercal chimaeras the whichLobe-finned fishes, Devonian,the during dominated are represented today a few only species of by and groups,huge and the whichspecies nearly into expanded have every niche.aquatic Osteichthyes Osteichthyes These are the of Theirabout with in skeleton 23,000 species. fishes, bony cludes nial bones supporting and the fins, Theare fins more refined ofthose than cartilaginous fishes, have jaws and the greaterthem allowing maneuverability, more freedom of Thin, movement. flexible noid protect and reducecus, the body drag water friction. An by to breathefishes these enables They when swimming. not also have a terminal mouth ( symmetrical is tail the of fishes, bony texture.The fallingteeth to be replacedout neweach Behind ones. by is a respiratory small eye the opening, have pairedfishes nous front of efficient an asymmetrical for swimming, the body), ( fins with ternalfertilization, They lackmammals. in as a der and internalfins Their olfactoryand senses developed. There are about 1000 recent species (and more forms)many fossil of cartilaginous includingfishes, two very differentgroups, more the primitive ( skates

, . ------rat (for (for skull plac semi , since meta spine, spine, or or limbs, or limbs, (a verti (a tetrapods paired olfac , migratinginto jaws inner ears chimaeras : , and the derived trait ). Their cartilaginous lateral line system pairedfins , or ) leading to to leading ) sharks anadromous cartilaginousray- fishes, (we have 3, used for balance). balance). have 3, for used (we , and paired surroundingand a the mouth nares and organ ( (deposit eggs). The remain young in secondary sucker , and a rudimentary are characterized by cartilaginous fishes . They include internal gills, to the , with spine-like processes. They also have eyes, Theyprocesses. spine-like , with also have eyes, cover the skin, protecting the skin, cover and giving a roughit and go to sea as adults for one or two more to sea for as adults and go and extrinsic (outside) eye muscles, two eye (outside) and extrinsic are respiration, used for rather feeding, than : Theseare fishes (smelling) (smelling) semicircular canal lens gills paired nostrils cally biting device biting made up ofcally elements endoskeletal or cartilaginousand a bony teeth), equipped with populations arepopulations decliningas a result of water pollution. Vertebrates Gnathostomata—Jawed Gnathostomes gill arches a with circular canals a special component ofear fishes which the inner in detects vibrationsand currents surroundingthe in water. the suckerthe to attach scraping and to other fishes skin their a raspingwith to sucktongue The blood. their sucker also is used to attach upstream make way their to stones as they Lampreys have a morespawning. during complete hagfishes’,than supports and skeletal called gills, the for morphose years, before returningto fresh They water spawning. for have a large, round on top of“nostril” single are head. the Many parasites, using however, and areclassifiednot sometimes as vertebrates. however, Lampreys freshwater to a larval stage of most to 7 years),life (up their then olfactory and perceptionbalance hearing, of of position movement) one with Their have a ventral they and heart. Hagfishesvertebrae,no have oid scales fishes, skates, raysfishes, skates, be a may skeletons arethey to have evolved thought from placoderm bony ancestors.called enameled scales tooth-like Small finned fishes, lobe-finned fishes lobe-finned fishes, finned Chondrichthyes Theseare the gill arches and tory organs 102 BIOL F150: Introduction to marine biology features of bony fishes are displayed bony fishes are by this huge group. of features advanced full suite of the and dinner table). The (and zonetothe ocean depths, in lakes and rivers the surf most fishes foundspecies.than 20,000 from include They osts Onegroup, the all vertebrates. of half for nearly fish diversity,and account 96%, about majorityof the vast called " by spines skin supported of webs finned bony fishes are Actinopterygii have sincecaughtin beentheIndian Ocean. more About 200 South Africa (C&Hp."A Fish Called Latimeria"). 171: coast of the found off was creatures thesestrange oneof when 1938, thought to beextinct it was until In fact, is an ancient form. coelacanth to the bloodbody andthelungs. flow The of and separation enameledteeth, appendages, they havefour similarly-sized of relatives living be the closest today,the most ancient fishes present to believed andare sorbs oxygen among and eliminates wastes. Lungfishes are ab bladder swim is a “lung” which and Africa.Their Australia speciesoccurin South America, today freshwater only afew but worldwide marinelungfishes occurred many speciesof the body on land. During the Devonian, used to support in the caseof which by fins containingpaddle-like bones andmuscles, rounded Sarcopterygii , is responsible for most of this diversity, with more for mostof , is responsible rays , the other type of living lobe-finned fish,also living the othertypeof , ." With." species, 23,000 they about makeup ( ( lobe-finned fishes ray-finned fishes ray-finned lungfishes tetrapods (and tetrapods) may be (andtetrapods) ): The fins of ray- fins of ): The ) are characterized characterized ) are . Like tetrapods, . Like tetrapods, tele - - Ray-finned fishes also include several small groups of of small groups several Ray-finned fishes also include by manygered otherfish species. human activities,as are endan Salmon populations are to spawnagain. survive often of the genusgives its name: including, changes in some cases,logical ahookednose which During spawning manythey anatomicalandphysio undergo tospawn and die. tothe natal stream precisely and returning Pacific in the North spending their adult lives streams, (birth) their natal miles from thousands of histories. may They travel and have extremely complex and sturgeons), life lampreys Salmon. PacificChum (Dog) salmon are on, Sockeye (Red) Salmon, and Salmon, Pink (Humpback) Pacific Salmon Pacific Halibut,Herring, Rockfish Alaska include around teleosts in waters Common as a cartilaginous “lungs.”skeleton, asymmetricaltail,or as acartilaginous such features, fishes tend to havesomeprimitive These fish. thanft long, 20 freshwater the largest more reach caviar. Pacific may Sturgeon pounds weigh and 400 The the Caspian Seais prized for its of Sturgeon Beluga The fin. andbow gars paddlefish, fishes, like the birchirs, water mostly fossils.” fresh “living are They bizarre-looking Atlantic Salmon Sturgeons are found in both fresh and salt water. found in both fresh are : Chinook (King) : (Silver) Salmon, Coho Salm , also is anadromous but these salmon , also is anadromous Oncorhynchus. Salmo Oncorhynchus. and five species of speciesof andfive anadromous , the genus, Pacific (like - - - - - Lesson 9: Marine Fishes 103 myomeres chromatophore iridophore and warning coloration and cryptic coloration disruptive coloration countershading and ovoviviparous, oviparous and viviparous anadromous and catadromous demersal and schooling secondary/derived trait inner ear, semicircular canal, , lateral inner ear, line and neuromasts olfactory organ/sac nares membranenictitating gill arch, gill filaments and lamellae erythrocyte, hemoglobin and myoglobin countercurrent system of flow • • • • • • • • • • • • • • • • • glossary terms glossary Salmo and , tail/caudal fin, anal fin, pectoral fins, pelvic fins bladder swim rectal gland and chloride cells Oncorhynchus placoid scale and cycloid/ctenoid scale spiracle and operculum heterocercaland homocercal tail Holocephali and Elasmobranchii Actinopterygii and Sarcopterygii ray-finned fish fish and lobe-finned teleost ostracoderm and placoderm and Gnathostomata Agnatha and Osteichthyes Chondrichthyes cartilaginous fish fish and bony vertebrae, backbone/spine/vertebral column and appendicular axial skeleton endoskeleton, skeleton, • • • • • • • • • • • • • • • • • your textbook. your Listed below areListed below terms some key Do a self-check this lesson. for understanding. for If you're unclear about the meaning of ofany these terms, re-read in this course the material and check textbook, guide and in your the glossary at the back of 104 BIOL F150: Introduction to marine biology online at eLearning.uaf.edu and in the course materialsyouonline ateLearning.uaf.edu received. andinthecourse a Include thesubject.) hasnoknowledge of someone who something,"describe" doso-don't just list. (To explaining that you to imagine are andshow your beclear understanding, own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 4. 3. 2. 1. ficiently. helpthemtoobtainoxygen fishes, which ef systemsof andrespiratory thecirculatory Describe four aspectsof andbony fishes, buoyancy intheways they andlocomotion. achieve betweensharks differences Discuss three infishes. coloration Describe four usesof Define, by thefollowing terms: comparingandcontrasting, n. m. l. k. j. i. h. g. f. e. d. c. b. a. cover sheet cover

, not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , chloride cells and rectal gland cellsandrectal chloride andcatadromous anadromous andovoviviparous oviparous fins finsanddorsal pectoral tail tailandhomocercal heterocercal (ctenoid)scales placoid scalesandcycloid andoperculum spiracle linesystem canalsandlateral semicircular Oncorhynchus andActinopterygii Sarcopterygii andElasmobranchs Holocephali Chondrichthyes andOsteichthyes Agnatha andGnathostomata andplacoderms ostracoderms Written Assignment for 9 Written Lesson Assignment and submit the written assignment to UAF eLearning & Distance Education according to directions found to directions & Distance Education according andsubmit the written assignment to UAF eLearning and (Please make your completeandconcise) answers Salmo • brief and clear , but, also complete . For example, if asked to asked For. example, if in yourin - Lesson 10: Marine Reptiles and birds 105 . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). found at the publisher's Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Know the distinguishing features of tetrapod the four groups and some of their evolutionary history. Be familiar with the marine representatives of reptiles are and birds, how they and some ofadapted a marine existence, for their ecological relationships. the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education. 277) and 12.13 (p. More information, including interactive flashcards, quizzes, videoclips can be and links, 1. 2. 1. 2. 3. and "Seabirds") 177–182 ("Marine Reptiles" and FiguresC&H: Chapter 12.12 9, pp. Reading assignment Reading steps for completing the lesson the completing steps for The first tetrapods were amphibian-like creatures four-limbed that emerged from the water onto land. All tetrapods descended from newly these terrestrial creatures. Thus, marine tetrapods have made an evolutionary round-trip—their ancestors came from onto land, and back the sea, to the sea. This lesson provides an overview of the tetrapods and an introduction to the marine reptiles and birds. this Lesson of Objectives and birds marine reptiles 10 106 BIOL F150: Introduction to marine biology Fig. 14 Classification Birds and of Reptiles Marine Kingdom Metazoa Kingdom EukaryaDomain Phylum Chordata

Subphylum Vertebrata Diapsida Anapsida Synapsida (mammals) Gnathostomata (jawed vertebrates) Sarcopterygii (lobe-finnedfishesandtetrapods) Sarcopterygii Tetrapoda Amniota Amphibia Birds: Order Gruiformes: rails, coots rails, Gruiformes: Order grebes Podicipediformes: Order loons Gaviiformes: Order bitterns egrets, herons, Ciconiiformes: Order geese ducks, Anseriformes: Order alcids, shorebirds skimmers, gulls, terns, Charadriiformes: Order frigatebirds pelicans, Pelecaniformes: boobies,Order cormorants, petrels shearwaters, fulmars, albatrosses, Porcellariiformes: Order penguins Sphenischiformes: Order crocodile saltwater Crocodilia: Order Squamata:seasnakes, marineiguana Order Chelonia:seaturtles Order Lesson 10: Marine Reptiles and birds 107

. - - - - am des and lungs . They . They and two 3-cham , unlike the , unlike ovovivipary Amniotes four fluid-filled four (jointed) bones to (jointed) scales ventricle calcification and to help prevent to help blood flow blood (to and from to conduct sound waves,sound to conduct , without a larval, without form. shell blood flow blood (to and from the cold-blooded ectothermscold-blooded (Fig 14). Amphibians include 14). (Fig Amphibians internal fertilization , with one , with articulated is one) surroundingone) is the embryo. systemic skin glands eardrums with dry with and skin pulmonary amniotes amnion can be relatively soft or hard, different . They have (the (the directdevelopment and the and the (“to creep”)(“to are (dryingout), and eyes adaptedof outside vision for and and eyes water, 3-chamberedheart with more with substantial, eggshells which separate the poikilotherms and soft permeableofskin a Most have amphibians. bered heart variousofmeans reproduction,including Their differenthaving species degrees of undergo sication eyelids limbs supporton land. Theweight their of name group this means have partially as they “double-life,” problem avoided the of returning by on land dessication waterto the part for of their life cycle. Their vulnerable eggs are usually water laid in and larvaethe and swim have gills about, wrigglingtails their areNo amphibians fish. like a lot and looking entirely marine. ancestorsAmphibian gave rise to the evolved a new type of egga with membranes These structures preventand provide dessication a stable gas and wastefor allow exchange environment, removal, the developing Livingand provide for embryo. amni food otes are classified commonly as reptiles, birds and mammals. Reptiles Tetrapods be dividedtwo general into may groups, the Tetrapods phibians frogs salamanders and toads, and newts. They have and a atria fromthe body) the Theylungs). also have ------• am . They evolved mammals marine reptiles andbirds and ancestorsto present-day akin lung fish . We have followed many branchings many have followed of . We

tetrapods lobe-finned the ofthe Tree here, Life to get from moving the Eukary n our survey of marine organisms, arrivedwe have at about remarkable these and the adaptations animals they basic requirementsthe have made to fulfill of their life in Thisenvironments. marine provideslesson overviewan of tetrapodsthe introductionan and reptilesmarine the to Marine mammals areand birds. covered lesson. next the in move, breathemove, keep and from dryingthe rawout—using material oflobe-finned fish ancestors. their Marine tetra turn,in pods, aquatic living, had to reinvent for themselves rawthe using of material land-adapted their ancestors. Keepevolutionary this perspective readmind, as you in trip. Their ancestorstrip. emerged fromsea onto land, and the tetrapodsall descended from newly these terrestrialcrea tures. Marine tetrapodsmade then a return back to a trip, Early existence. marine tetrapods adaptednew to the chal oflenges livinghow to support on land—like themselves, mediate between lobe-finned fishes and tetrapods,mediate between lobe-finned fishes which representmay two groups,between the a link was recently 80: SteppedFishes “When on Land”). discovered (C&H p. Thus,tetrapods marine have made an evolutionary round- with one upper bone, two lower bones and many small bones, bones, small bones and many lower upper bone, two one with bones and other Skull and foot. ankle the leg, in those like parts ofalso are the skeleton some lobe-finned in similar of and tetrapods.fishes Fossils featureswith a fish inter phibians, reptiles, birds from probablyaround Devonianthe during fishes, 350–370 mya supports bony some lobe-finned In fishes, Period. the fins in are arranged muchoflimbs the bones in the like tetrapods, ote branch, to deuterostomes, to chordates, to animals, to vertebrates, to jawed vertebrates, to sarcopterygians, and include ("four-footed") to tetrapods.finally Tetrapods I 108 BIOL F150: Introduction to marine biology and flying forms. They are still the most diverse amniotes, still the mostdiverse are They and flying forms. swimming with slithering,running, crawling, ingly diverse, flexible for amaz feeding.more were In the past, diapsids many fusedboneswith openings, making them lighter and skulls include diapsids. Their are plus the birds, reptiles, All the other diverse. very previously were anapsids saurs, thedino of the age in Dating solid from bone. covered completely skulls are whose theanapsids, of representatives the only living are reptiles, the most primitive Turtles, ture. Diapsida assigned tothegroups, these animals. are They of themarineforms to classify 14 usedin Figure mals was andmam birds forreptiles, scheme A commonclassification inside the mother. asit develops wastes, andremoving bryo into the mammals are lay but (the platypus and echidna) eggs, all other tremes Mono spatial reasoning). motor coordination, perception, neocortex called the with a unique region brain also have a large They their with a Mammals rate. tosustain a higher pods,metabolic andallows birds than the tionally the long bones, allowing air tobecirculated mented by a complex system of blood flow. supple lungs are temic andpulmonary Their sys of and completeseparation circulation efficient more heart 4-chambered a evolved fusedorlost in birds. They Many bonesare sternum light yet strong,and in some caseshollow,with a and hair), and abony skin of cells and made from into fashioned are with Birds include locomotion, and manyReptiles have evolved modes of hard shells hard hair are are 4-chambered heart 4-chambered placenta turtles, snakes, lizards turtles, andcontinuously. system efficient is a more This tidal flow (breastbone) to support powerful flight muscles. powerfulflight tosupport (breastbone) and or , like birds, are are , like birds, endotherms , which is involved in higher functions (sensory is involved in higher functions (sensory , which viviparous fur Synapsida , and milk-producing and milk-producing , of of and wings (inhalation-exhalation) of othertetra (inhalation-exhalation) of calcium carbonatecalcium with two atria and two ventricles for with two atriaandventricles beak umbilical cord umbilical and endotherms . The egg membranes are modified are membranes egg The . , based partly ontheir skull struc partly , based . They have . They without teeth. Their bones are bones are without teeth. Their homeotherms . They are distinguished by are . They keratin air sacs , and , (like our fingernails (like our fingernails mammary glands mammary and , nourishing the em feathers . Their forelimbs forelimbs . Their homeotherms extending into crocodilians that lay eggs that lay eggs Anapsida, unidirec derived derived keeled ------. .

tral tral adorsal ized their shell, made upof Chelonia Crocodilia lonia majororders: the three of representatives clude to the sea, but these in species have returned ful of Only ahand species distributed 7,000 worldwide. about with vertebrates, successful terrestrial are ing reptiles diversity, their former the liv shadow of Although a mere Marine Reptiles Period, the land during the Permian the synapsids. ruled animals that reptile-like of group diverse a descendants of the onlywith species.over 14,600 surviving Mammalsare iguana snakes species of 55 about including marine reptiles, numerous alsothemost are They reptiles. living of anddiverse merous Squamata (C&H p.180:Sea Turtles”). carelessness Endangered “The or considered are All turtles sea rine predators. and ma for terrestrial easyprey are hatchlings ians! The amphib miles,thus leading adouble lifeoppositetothat of than a thousand more lay sometimes traveling their eggs, to toshore still return forswimming. Female turtles sea turtles Onlyall continents 9species—the except Antarctica. species, found on terrestrial than 200 more are jaws. There down with teeth,butpowerful can clamp lack Turtles cage. in a rigid rib enclosed are the lungs,and deflating which that aid in inflating alsohave special muscles They tracts). and reproductive (common exit forand cloaca the excretory get turtles their air through they breathe Like all tetrapods, acids accumulate in the blood. metabolic when bernation, and for thebloodbuffering forduring hi makingeggshells calcium can bewithdrawn which as a“calcium bank” from the shell serves In addition to protection, their ancestors. of the shoulder andhip girdles and elements of tebrae endangered plastron (turtles), (turtles), . The elongate, limbless snake body is pre-adapted limbless snakebody elongate, is pre-adapted The . —are marine, with marine, into limbs modified —are found mostly in tropical waters, and the found waters, mostly in tropical ( ( (crocodiles, alligators, caymans andgavial). alligators, (crocodiles, additional gas exchange gas additional turtles snakes . The shell developed from scales, ribs,from ver shell developed . The , the victims of human and use, abuse the victims , of Squamata ): These ancient reptiles are character are ancient reptiles ): These and lizards (snakes and lizards), and (snakes and lizards), ): These are the most nu are ): These carapace through the throat the throat through threatened lungs andven flippers marine Che , but sea sea ------

Lesson 10: Marine Reptiles and birds 109

------Cor . Terns , which plum webbed Homeo use them use use their down located near . All birds. All nest . Theyto live tend for waterproofing for . Thein salt excess albatrosses shearwaters salt glands shorebirds colonies saltwatercrocodile and oil glands and seabirds convergent evolution puffins, murres : Thiscategory includesa number of unrelated areTheseexception. an birdsancient lackoil the and the abilityand the birdsallowed fly have to to invade . Some use them for propulsion for . Some use them when or paddling diving. of seabirds restrictedmostly is to black, whiteand grey back on the body than in land birds,land backin a more than body the on efficientposi Most seabirds swimmingfor and diving. tion have feet Others, like as stabilizers whilediving, during webbed feet to provideleverage whenoffpushing against Most seabirdsin nest water. the more and invest time have fewer young, breed later, longer, the ocean, except when breeding. Shorebirds, on the other and estuaries to stickhand, tend bays, coast, using to the migration. during especially feedingand shelter, mudflats for Seabirds sharethey groups, some adaptationsbut to a marine lifestyle, as a resultof seabirds by extracted is gested by and excreted through cavity or nasal mouth. the the eyes, have other birds,Like they seabirds but plumage, their are more protected from the denser feathers ofwet and cold by and a layer morants gland responsible waterproofing, for reducing buoyancy their to divethem and allowing deepersearch in of food. also lack waterproof catch they plumage, since prey their at Theor above surface, the rarely touchingwater. the age colors, to provide camouflage. Their legs are set farther ing on all four legs. They legs. arefour all largeon their ing oviparous, laying eggsThe land. on females ofprotect species some nest the among reptiles.behavior a unique careand young, the for species occurs,One marine the reachesoflengths coastal watersin 20–30 found ft, and is aroundAustralia, and some western Indonesia islands. Pacific Birds Marine ofdescendants Also “ruling the reptiles,” birds aremost the diverse tetrapods, about with 10,000 species. thermy Birdsrepresent marine environments in environments. many 9 ofout about 30 orders of 9.1), and about birds Fig. (C&H 3% of Theybird all are species. often dividedtwo unof into categories, ficial seabirds land, but on spend muchof lives their or above on

, ------Salt- behav to subdue salt-excreting salt-excreting venom 4-chambered heart enhanced gasenhanced exchange to assist with breathing,with to assist and stretches oflength the almost (bony separation (bony of and the mouth located above to help them the nostrils diaphragm right lung inhabit spend the Galapagosand Islands, : These ponderous, armoredof descendants by flatteningbodies againsttheir by warmthe rocksur that sits under the tongue. Like their terrestrialtheir Like tongue. the under sits that coun secondary palate quickly on land by belly-crawling, walkingand even quicklyby on land gallop submerged with only the nares above water. They have well- nares the submergedonly with above water. developedareand senses efficientpredators, conical with They massive their use powerful jaws. teeth set in for tail and can move swimmingand to sweep preydeep into water, ous crocodilianshave a complicated (capable ofa 3-chamberedlike functioning heart during dives), a muscular a whichcavities) nasal breathing to continue them allows while tected in the Galapagos Islands, a national park a national Galapagos the tected in Islands, of Ecuador. Crocodilia the once “rulingreptiles” are some endowed surpriswith advancedingly features. other reptiles, Unlike the amphibi iorally elevatingface, and cool down by torsothe to it and exposing breeze,the cooling opening and closingthe in vessels blood chestto control They exchange. heat can also pant to cool fewhave Marine iguanas natural off. predators and are pro ft and remaining underwater as long as 30 minutes. for glands excreting ofcontent salt high the cope with diet. They their sneeze also whichsalt, frequentlyexpell to gives blacktheir bodies a whitewashedwarm look. Marine iguanas themselves Marine iguanas divingthe cold water into their days to graze on seaweed, warmthe on and basking rocks. Theyto swimtails their use claws, their diving and clingas deep bottom by as 40 to the their prey. Most sea snakes are Most sea snakes entirelyovovi and marine prey. their returnsome species still viparous, but eggs. to land to lay areSome species becoming rare, are as they for hunted and areskins their inadvertentlytrawlin caught nets. fishing as they are able to absorb additional oxygen through the areas they to absorb able oxygen additional and throughskin, swallowingthroat the by water and then a by eliminated is salt Excess it. ejecting gland terparts, are they carnivorousand use for efficientimprovedhave snakes and sea for swimming, on laterally plan, with body this flattened bodies paddle- and Theshaped tails. stores when increasingunderwater. oxygen their body, the Like turtles, sea snakes have 110 BIOL F150: Introduction to marine biology surface, using a variety of of using variety a surface, belowthe prey Other speciescatch andsometerns). birds mers), on the wing,prey by while krill, fishes and squids. Some as such prey may concentrate oceancurrents where face, and fill different feeding strategies species to adoptdifferent allows different theirwings,and body,beak feet, modifications of of A variety in length and width. intermediate gulls are wings of purpose all- pointed wingsanddipping,for the more swooping while have dives. Terns or deep flights and jaegers direct short, them in propel their body weight), which of ing for one-third (account muscles breast wings with massive shorter much have gliding. Murres effortless ft) for long-distance, nearly have long,slender powerfulwings (wingspans can exceed7 in their different is reflected which in the specific ways they also differ fly,Seabirds or dive, swim and winter in the Antarctic! in the Arctic miles to breed 25,000 their traveling young. Tern Arctic rear holds the record, The migrate skuas. Somespecies inland to nest, like the tundra-nesting Some speciesmove pelicans, and frigatebirds. cormorants, the oceanforfood. coastal,but on still dependent strongly more are seabirds the of (members murres and and petrels, ists into divided for food. mayPelagic be further seabirds tering on orin the open ocean,totally on the sea dependent are in how they greatly use the ocean. Some also vary seabirds due toconvergent evolution, sometraits While they share tolife on land. compared predators of lack but also a relative lifeatsea, of tothechallenges may in response haveevolved ten ods away, of are involved usually andpairs are both parents longperi thenest site andrequire mayfrom takeplacefar for foodinvestment, andbecauseforaging this greater of them, than do other birds. Because into raising and energy monogamous pelagic like the swimming pattering , as far as thousands of miles each year, miles each asthousands tonest of and , asfar , breeding on sea cliffs andsmall islands and win onseacliffs , breeding tubenoses niches open-water swimmers open-water (e.g., shearwaters). Others catch their catch Others (e.g., shearwaters). (storm-petrels), or (storm-petrels), Coastal . The reproductive strategies of seabirds seabirds of strategies reproductive The . dipping . Many birds feed at the water sur at the feed water . Many birds (albatrosses, shearwaters, fulmars fulmars shearwaters, (albatrosses, alcid seabirds include gulls and terns, gulls and terns, include seabirds diving surface feeders surface family) and penguins. Other (gulls), wing types styles, and still others styles,and still others snatching skimming like puffins and like puffins . Albatrosses . Albatrosses catch prey prey catch (frigate aerial (skim ------in guiding fishermen to fish, and sailors to land. The rela toland.The tofish,andsailors in guiding fishermen role alsohaveplayedsome controls.Seabirds a traditional continues today,egging although less intensely andwith Hunting and Auk. the Great the completeextinctioneven of populations,caused local or of to thedecline In many casesthis has contributedfood, oil and feathers. for andhunted birds eggs seabird people haveharvested their populations in manytinue to affect ways. Traditionally, andcon Humans have hadalong association with seabirds, called is This resources. the available of parts utilizing different specialization, species coexist through in thesamehabitat on small adult birds. In this way, oreven seabirds, multiple enge Skuas, otherseabirds. gulls andgiant petrels foodof in engage on prey at different depths within the sediment (C&H Fig. depths at different on prey to beaklength, feeding according divided further are ers gleaning their resources. partition and niches allow them toexploit various which ing strategies feed diverse haveevolved shorebirds Like seabirds, mph. upto50 miles, as long as15,000 atspeedsof migrations them on pointedwings carry long, narrow relatively Their andfishes in the sediment or shallow water. invertebrates in hunting or coastal wetlands,for while small along beaches forfeet, wading without webbed pointed beaksandlong legs mostly small with stilts andavosets. are They plovers, ers, sandpip include skuas and thealcids.“True” shorebirds iformes Shorebirds these losses. someof helping to reduce are tion measures drift nets bird of establishment cats. The and feral species, like rats structive de oil spills, toxins of and pollution, and the introduction of fisheries (C&H Fig.p. 17.6, victims alsoare Seabirds 389). ment year, each albatrosses 100,000 of asaresult anestimated is the loss of One serious problem affected. negatively are butis complicated, in many casesseabirds today andthe fishing industry tionship between seabirds dead animals and prey on the eggs or chicks of other of chicks or ontheeggs animals dead and prey in nets set out for and swordfish by in nets set out for longline tuna and swordfish resource partitioning resource sanctuaries , which also includes some seabirds like gulls, terns, like gulls,terns, seabirds some alsoincludes which , at the surface are two general strategies. Prob strategies. two general are atthe surface aerial pursuit , and other fisheries regulations and conserva , andotherfisheries regulations : Shorebirds belong to the Order belongtotheOrder Shorebirds : and Probing refuges . Frigatebirds andskuas steal the Frigatebirds . (C&HFigs.and 9.8.). 9.7 into mud, and the sandor , an international ban on , aninternational extinctions entangle Charadri scav ------, 111 Lesson 10: Marine Reptiles and birds

- - - - or of , and of 1929. threatened Audubon Society Audubon Endangered Species Act Endangered Endangered Species Preservation Endangered Federal Migratory Act Bird Treaty Migratory Bird Conservation Act . The of the by 1966, followed their eleganttheir feathers,of millions herons egrets and were late 1800s. the in fashion the eachkilled industry, for year Thisprovoked first the and action, environmental public resultedof foundation the in the passage of the Migratory are animals particularlyvulnerable to disturbances suchwhichand pollution, spills oil as have destructive ef shorebird popula Sadly, fects on habitatswebs. and food develophuman continued with aretions declining still today, ment of species are Many coastal habitats. 1918 the and endangered Act of Fish U.S. and the 1973, which NOAA administeredis by Service, and Wildlife were enacted to protectendan both gered species and the ecosystems upon which depend. they - - - as herons,as egrets, bitternsrails,and swimmers and divand ers grebes loons, Prized for ducks, like geese, and coots. Members of five other orders of birds, besides Charadri iformes, areofto those similar ways sea in to the connected “true” shorebirds. These include long-legged waders such food. Other shorebirdsfood. have evolvedstrategies, unique e.g., oystercatchersof open the shells and clams mussels using thicktheir and turnstonesbill, triangular wander beach, the turning stones and debris, searchin over of choice mor Thesesels. are of examples good all resource partitioning. 12.12, p. 277). The has example, Curlew, Long-billed for 12.12, p. 9 inches),to beak (up and probeslongest the deeper than the relatively short-billedGleaners Willet. also divide their habitataccording leg to wading length, differentdistances surfthe into scurrythey as beach the along searchin of 112 BIOL F150: Introduction to marine biology your textbook. of attheback theglossary guideandinyour textbook, andcheck thematerialinthiscourse re-read theseterms, any of themeaningof about unclear you're If for understanding. for thislesson.Doaself-check somekey terms Listed beloware • • • • • • • • • • • • • tidal flow and unidirectional flow tidal flowandunidirectional blood flow systemic andpulmonary atria andventricles andhomeotherm poikilotherm andendotherm ectotherm amnion alcidsandtubenoses Charadriiformes, andshorebird seabird Chelonia, Squamata andCrocodilia Anapsida, DiapsidaandSynapsida Reptilia, Aves andMammalia amphibian andamniote tetrapod glossary terms • • • • • • • • • • • diaphragm air sacs salt glandandoil keratin andcalciumcarbonate andplastron carapace ing partition andresource evolution convergent andviviparous ovoviparous oviparous, development direct threatened andendangered species guano cloaca - 113 Lesson 10: Marine Reptiles and birds in your in your . For example, if example, . For asked to complete , but also , but clear and brief • (Please make your answers and concise) complete your make (Please and submit the written assignment to UAF eLearning to UAF assignment written the and submit according Education & Distance to directions found Written Assignment LessonWritten 1o for carapace and feathers seabird and shorebird tubenoses and alcids oil glands and salt glands pulmonary flow and systemic blood evolutionconvergent and resource partitioning threatened and endangered species amniote and amphibian anapsid and diapsid poikilotherm and homeotherm Charadriiformes and Chelonia , ofnot those or Course textbook the Guide. Be

cover sheet k. g. h. i. j. c. d. e. f. a. b. Describe an example ofDescribe an example learned evolution convergent that you about in this lesson. Compare adaptations gas in sea turtles, for exchange birds. sea snakes and Discuss some of the differences in feeding strategies among birds (choose either shorebirds or seabirds), which result in different niches different for allowing them to coexist. species, Tetrapoda means four-footed. Describe how the limbs of means four-footed. Tetrapoda sea turtles, and how they snakes and seabirds sea differ, are adapted to the lifestyles of these groups. Describe the differences between the eggs of reptiles amphibians, and birds. Define, by comparing and contrasting, terms:the following Define, by 6. 4. 5. 2. 3. 1. online at eLearning.uaf.edu and in the course received.online at eLearning.uaf.edu materials you own words understanding, be clearyour and show are imagineto you that explaining (To list. just do so - don't "describe" something, someone who of has no knowledge the subject.) Includea Study the information presented for this lesson until you feel that you have a good understanding. Write your answers your information the understanding. have a good Study Write feel that you presented you until lesson this for 114 BIOL F150: Introduction to marine biology 115 Lesson 11: Marine Mammals - Marine

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). found at the publisher's Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Be familiar with the various groups of their distinguishing characmammals, marine teristics and representative species. Understand adapted how marine mammals are for and physiologically anatomically life in the sea. the information of pages Read this lesson in the following for this Course Guide. Complete the Biology Complete the eLearning & Distance Education. 1. 2. 1. 2. 3. 186–209 ("Marine Mammals" ) C&H: Chapter 9, pp. More information, including interactive flashcards, quizzes, videoclips can be and links, Reading assignment Reading steps for completing the lesson the completing steps for In this lesson we conclude our survey of marine organisms animals with marine mammals, so well-adapted life in the ocean that it is difficult are for to believe they members of our own class. this Lesson of Objectives mammals marine 11 116 BIOL F150: Introduction to marine biology Kingdom Metazoa Kingdom EukaryaDomain Fig. 15Classification of mammals Marine Phylum Chordata SubphylumVertebrata Family Family Phocoenidae: Monodontidae: porpoises Suborder Beluga, (6 Family Odontoceti—toothed Narwhal Family species) Neobalaenidae: whales Family Eschrichtiidae: Pygmy Balaenopteridae: Gray Suborder Right rorquals Whale Mysticeti—baleen Whale (10 Family whales Family species) Trichechidae: Dugongidae: manatees Dugong (3 species) Family Odobenidae: Family Walrus Family Mustelidae: Ursidae: marine Polar otters Bear Subclass Eutheria/Placentalia—placental (Subclass mammals Prototheria/Monotremata—egg-laying Amniota mammals) Gnathostomata (jawed vertebrates)

Sarcopterygii (lobe-finnedfishesandtetrapods) Sarcopterygii Tetrapoda Class Mammalia (Subclass Metatheria/Marsupialia—opossums, kangaroos, etc.) kangaroos, Metatheria/Marsupialia—opossums, (Subclass Superorder Ungulata (horses, elephants, deer, Ungulata(horses, Superorder cattle,etc.) Order Order Cetacea Order Sirenia Order Carnivora Suborder Caniformia—"dog-like" (dogs, skunks, racoons, etc.) (dogs, skunks, racoons, Caniformia—"dog-like" Suborder Feliformia—"cat-like"—cats, civits, hyenas,(Suborder etc.) Superfamily Pinnipedia (Superfamily Platanistoidea: river dolphins) Family Physeteridae: sperm whales (3species) Family whales Physeteridae: sperm Family Delphinidae: dolphins,(37species) killerwhales Family (4species) rightwhales Balaenidae: Bowhead, seals(19species) Family Phocidae:true Family Otariidae:sea lions, furseals(14species) Family (21species) Ziphidae:beakedwhales 117 Lesson 11: Marine Mammals - - - - . They can con ," because ribs their collapsible (without oxygen). These adapta oxygen). (without thermoregulation , in which, in heart rate dramatically slows bends ) are) of other examples extraordinary the ) and blood flowreis tissues to nonessential blood and ) anaerobically diving response echolocation bradycardia adaptations mammals have made the sea. marine to life in classificationA Figurein schemeshown is mammals marine for 15. Once again, begins it and summarizes Domain their with major branchinglong evolutionaryalong the points by paths which came to be. animals these Thisscheme differs from arepinnipeds 9.1), that in (Fig. uses textbook the one your includedOrderin Carnivora, ofanother example just science is maintained, and blood oxygen stores muchlast (in longer oxygen and blood maintained, is more for some cases, an hour). Thethan diving response of marine mammals (and other divinga highly is animals) developed form of a basic protective response all shared by vertebrates.divecan also mammals marine Some to great depths(more 7,000 than ft, the case ofin sperm whales) the " getting without absorptionthe limit and lungs of nitrogen blood. the into The is used abilityalso control to selectively distribution blood marine mammals for by serveextremitiesthe heat in or dissipate (even tongue, the in case ofthe in great the whales),as needed. The ability to and waterutilize present inspiredfood, in air and blubber, marine enables to excreteseawater, than saltier is that urine acute tactile hearing, Excellent mammals to conserve water. ability the underwater to communicate senses, sound, with ofuse and the waves sound to locate, and perhapsprey stun, ( and use it more efficiently, and their tissues aretissues their and able better moreit use and efficiently, to function periods underwa mammals to spend long marine allow tions their breath. holding Breath-holdingmarine mammals in ter, adaptation a physiological called greatlyis by also extended the ( duced. Blood flowbrainthe like tissues to essential heartand

, . ------• (at mam fur flippers , nourish tetrapods or hair marine mammals marine mammals with . All marine mammals marine . All and bearing live young. and bearing live young. , all of, all whichgives a small them . In most, the externalmost, the . In ears are placental mammals placental placenta neocortex

blubber . They areevery in found ocean and on

that produce milk for their young, and a produce that young, their for milk

. These anatomical featuresheat help to minimize

polar bearspolar air-breathing homeotherms and ofmarine organisms, all with made the return-tripMammals have independently to n this lesson we conclude we lesson n this our survey of the

flukes

cetaceans, sirenians,cetaceans, pinnipeds, marine otters Marine mammals also have made extraordinaryalso Marine mammals physiological adaptationsocean. the Theirto life in grow more young quick a fatwith on milk as 40–50% as high milk content (human ly store more3%is fat). oxygen They breathe more efficiently, small or small absent, and externalare and nipples genitalia hidden or depressions.slits in Theare limbs absent or partly con and are greatly modified into body, the within tained and streamliningmaximize and loss, and propulsion water. the in Marine mammals are large (including the largestever animal inhabitto Earth,the Whale, growingBlue the 100 ft over to towardand tend and 150 tons) well- being spherical, the fatendowed with or surface-to-volume ratio large brainincluding a belong to the subclassof the embryoing through a They are long-lived, reproductiveand have a slow rate.. membersof they mammals, all like But, class. our own are life, even during some time if a few just whiskers), mary glands everya varietyin continent, of ecological roles, includ herbivoresing (sirenians),filter feeders whales), (baleen carnivores carnivores and top (pinnipeds) whales (killer and polar bears). are they some cases, so well-adapt In difficultis it ocean that the life in areto believe ed for they a marine existence at least five times, with the evolutionthe with five at least times, existence a marine of and I 118 BIOL F150: Introduction to marine biology threatened marine pollution, oil spills in particular. Many speciesare by all types of Cetaceans may operations. beharmed val during na have beenlinked to high intensity sonar bursts water, for ship propellers. example from to noise “pollution” in the to sound makes them vulnerable cetaceans of sensitivity mines. underwater The and mark by to detect theNavy trained are bottlenose dolphins,which sophisticated of capable Some odontocetesalso are their direction. discern vocalizations well known fortheir are They rate. with low reproductive a highly long-lived, intelligent socialanimals are Cetaceans prey, larger or grasp teeth used tostrain usually fish or squid. Odontocetes have peg-like benthic feeders. are which whales, krill and small the fish, except gray baleen plates, trapping their through water es). Mysticetes filter huge mouthfuls of toothed whales and whales) and gray (e.g., whales bowhead two groups, Cetaceans include totwohours. for close submerged overft andremain 7,000 to can dive mph, others at 25 swimming of capable ceans are a forming their head, on top of are layer of sal fin zontal with ers, and div the most specialized mammalian swimmers are They theiryoung andsuckle in the water.birth give mate, feed, ago.million 50 years Cetaceans to theseaabout returning animals like began cattle anddeer) lated tocloven-hoofed (surprisingly,pletely ancestors aquatic mammals. Their re all com “ketos” meaning“seamonster”) are Greek (from dolphins Cetaceans Order Cetacea below, detail in your textbook. discussed in greater andare Sirenia the orders of all members including groups, marine mammal as an ongoing The quest, subject to debate. flukes . Their highly compressed neck vertebrae and a thick and a thick vertebrae neck highly compressed . Their , and some members of of members and some , blubber flippers and : The approximately 90 species of approximatelyspecies of 90 : The or that propel with a vertical motion, and a with a vertical that propel , and can hear sounds manyand canhear , miles away,and porpoises echolocation add to their streamlining, andtheir nostrils addtotheir streamlining, endangered (e.g., sperm whales, dolphins and porpois whales, (e.g.,sperm for forelimbs, no external hindlimbs, hori for noexternal forelimbs, who comprise the Order Cetacea comprisethe Order who Mysticeti , primarily because of over- primarily, of because , perhaps best developed in bestdeveloped , perhaps Carnivora blowhole or Stranding baleen whales , are introduced introduced are , Odontoceti Cetacea . Someceta whales, whales, deaths dor and or ------

tolerate cold water despite their large sizeand despitetheir large coldwater tolerate they with Slow,not a lowmetabolism, do creatures passive herbivores gent evolution many (another example similaradaptations of share lineage is as oldcetaceans',andthey mammals. Their of the only aquaticgroup otherentirely are sirenians mermaids, Sirenians Order Sirenia have helpedsometorecover, continue. practices but harmful populations whale and conserve human activities regulate Act Protection Mammal mission the gear. of formation The dolphins in fishing and entanglement of whales hunting of About 35 species belong to three families: families: species belongtothree About 35 coaststo the openocean. and from tothe tropics, packs ice the polar from ily range fish,shellfish and squid.They All are water). in deep, murky by theNavytolocateobjects (sea lions have beentrained to function both in air andin water adapted organs sory flippers webbed with four large, and rather sleek-bodied mammals. are They andwith other marine to bears), related most closely are (they relatives carnivore teristics with both their terrestrial somecharac Pinnipeds share anddivers. swimmers perb young, orto their andsuckle birth give onto theicetomate, or ashore come ties with some land. They (“wingedfeet”) still retain pinnipeds to the sea than cetaceans or sirenians, returned Pinnipeds Order Carnivora Cow A fifthspecies, the and coastaldevelopment. tohuman population growth boats andotherimpacts related loss,the past, and today habitat pollution, collisions with face andallare remain, called Commonly etation, alongcoastsandin estuaries and rivers. with adequate veg shallow waters to warm, restricted are , was hunted to extinction as recently as about 1770. hunted to extinction asabout , was as recently manatees, dugongs in 1946, passage of the United States of passage in 1946, : So named because they were oncemistaken for: So named becausethey were ( , like their closest living relatives, theelephants. relatives, living , like their closest seals, sealions molt ). Unlike ). other marinemammals,they are . They are awkward on land, but are su onland,but are awkward are . They , have both , endangered of 1972, and otherattemptsto 1972, of International Whaling Com Whaling International or carnivorous and fur sea cows . They were slaughtered in slaughtered were They . and walrus blubber , only four species ): More recently recently ): More , eating primar blubber Steller’s Sea Steller’s Otariidae , and sen , conver Marine . They They . ------

Lesson 11: Marine Mammals 119 , ------, polar polynas in 2008. in , and a layer of, and a layer (whichalso depend threatened ” when scarce.is food circumpolar Arctic guard hairs ringed seals ringed . Naturalalso become substances con . Theand thickness extent of the sea ice in walking hibernation on land or on packor on land on ice, where give birth. they dens Thedeterioration of correlatedis sea ice a decline with in and reproductive the physical of health polar bears. Loss of habitat, their and their declining reand slow populations productive rate, whichabilitytheir limit to replace individu areals, grave cause for concernfuture the for ofspe this were they as listed the U.S., In cies. tributing their weight on the ice, like snowshoes. They aresnowshoes. ice, like the on weight their tributing extremelyequipped stout, curvedwith claws clambering for bears Polar have an extraor ice and gripping prey. onto the dinary or water, food periods without long for ability go to a “ entering Circumpolarpolar peoples bears, have long hunted and there has been concern about over-harvestingof spe this the top of At cies. trophic the also arepyramid, they espe vulnerablecially which to pollutants, become concentrated eachwith chain,step food the up called a phenomenon biomagnification centrated, polar bear liverand eating "hyper-vi cause can levels polar of due to toxic A" taminosis A. However, Vitamin unprecedentedbears threat, with face now an even greater, rapidthe deteriorationof sea-ice habitat their as a result of change climate and it has been breaking Arctic has decreased drastically, up earlierrecent in we learnedas decades, Lesson 3. in the ice-covered waters of the bears dependsea ice as a platform on travel for and ac hunt breeding,mates and seeking and for for cess to land, ing their primary prey, on sea ice). Theyare particularly dependent on areas ofopen water surrounded3), which(Lesson ice by are richly often productive and attractive to marine mam and seabirds.mals travel They may on distances long search in of water, the or in ice the Males or mates. food out on the packoverwinter ice, whilepregnant females dig bearshuge these up to 1,700 pounds, have a Weighing shortears,and small snout and are covereddense with whitewater-repellentfur with fat to 5 inches up thick. Theirfeet also are and fur huge coveredoarsas bottom, acting the on waterthe in dis and

. . ------Odo groom ) and endangered endangered webbed hindfeet and earless seals tail , whichSouth the in found is ( , whichinhabits Norththe Pacific ). Also over-hunted for their skin, their for over-hunted ). Also Phocidae ), and the smallest marine mammals. At marine mammals. smallest ), and the : These members of the Carnivora or chungungo ), sea otter : Thelargest carnivorousand most bear (fam ), the polar bear is the marine mammal least polar bearthe ), the is the walrus ( as its primary as its throughout habitat. Distributed Mustelidae . Demand for pelt nearlyits . Demand for led to the sea otter’s ex ofair providedis insulation mammal. Additional by any Ursidae eared seals Polar Bear Polar ily relianttotally is on adapted yet existence, to an aquatic sea ice benthic invertebratesbenthic urchinsclams, like mussels, and crabs. One ofbesides primates known animals only the rocks use they tools, to use to pryor shells prey their offrocks, the and to hammer or pryshells. open their Sea otters are oil to disease, habitat also subject loss, trash,plastic in entanglement spills, and conflictabawith ranged they as far Baja as south Previously, lone fisheries. wacoastal the in only California,be found can today but tersof northern the Sea ottersPacific. on feed primarily trapped in the fur, whichbe must kept clean their sur for trapped the fur, in vival, so sea ottersspend about 10% oftime their ing remainsspecies the and today tinction, used for swimmingrelatively and making used for short, shallow Theirdives. front paws lack webbing and have retractable Theyclaws groomingblubber, have no for used and feeding. a relativelyhave but metabolic high rate, densest and the fur rest, mate, give birthwater. the in and suckle young their relativeAs newcomersottersat sea, marine to life lack ofmany adaptations the mammals. other marine seen in The flat a long, sea otter has Ocean and the Ocean, areeast Pacific otters only the feed that exclusively ashore go often prey, at sea. to eat their Chungungos rest, give birth whereasrearand young, their sea otters der arelargest the representatives offamweasel the ( ily 6 least ofotter species are or par 13 the existing fully They adapted at high occur all existence. to a marine tially Thelatitudes. ( benidae remain species many blubber, meat and Marine Otters 120 BIOL F150: Introduction to marine biology your textbook. of attheback theglossary guideandinyour textbook, andcheck thematerialinthiscourse re-read theseterms, any of themeaningof about unclear you're If for understanding. for thislesson.Doaself-check somekey terms Listed beloware • • • • • • • • • • • • • • • • • baleen plate andspout/blowblowhole and flukes flippers fur, andblubber hairs guard neocortex andplacenta uterus glandsandmilk mammary andseaotter marine otters Mustelidae andUrsidae sealsandwalrus seals,earless eared Otariidae, PhocidaeandOdobenidae manatees, seacowsanddugongs andgreat whales rorquals andtoothedwhales baleen whales dolphinsandporpoises whales, Mysticetes andOdontocetes dia) (andPinnipe andCarnivora Cetacea, Sirenia glossary terms - • • • • • • • • • • • • • • • • lactic acid the bends apneustic breathing bradycardia/bradychardia response diving thermoregulation andendotherm homeotherm viviparous andambergris organ melon, spermaceti Mammal Protection Act Mammal Protection CommissionandMarine Whaling International delayed implantation migrations stranding/beaching breaching pods vocalizations andecholocation 121 Lesson 11: Marine Mammals - - - in your in your . For example, if example, . For asked to complete , but also , but clear and brief Allow 4 hours exam. for completing the (Please make your answers and concise) complete your make (Please and submit the written assignment to UAF eLearning to UAF assignment written the and submit according Education & Distance to directions found Written Assignment LessonWritten 11 for (If you live in the Fairbanks area there is no need to complete the form. Come to our office during our office Come to the form. complete need to is no there area Fairbanks in the (If you live , ofnot those or Course textbook the Guide. Be cover sheet What reasons think of can you there for endangered being so many and threatened species of marine mammals? Briefly describe the feeding habitsBriefly of one species of marine mammal. Describe one unique behavior of a marine mammal that interests you. ofDescribe one example unrelated evolution convergent involving of species Describe another marine mammals. ofexample marine mammals and some non-mammalian animals. evolution, convergent involving following are most closely related: sea squirts,following insects, you octopuses, jellyfishes? Like land mammals (and other tetrapods), marine mammals are air-breathers. adap physiological Describe four tations of the respiratory systems of that increase marine mammals the efficiency of gas in a marine exchange environment. for minimizing heat loss in marine mammals. for List the three families describe two distinguishing characteristics of and one repre each and, for pinnipeds family, (again,sentative species. the name of a unique organism) which which of of seal? To To ringed dugong, is a sea otter most closely the related:following the walrus, polar bear, Name three orders of ("species" mammals that include and two marine species from marine mammals, each order. are unique organisms "whale" example, not a species) is - for adaptations anatomical Describe four fromHeat is conducted away warm bodies 23 times faster in water than in air. 9. 6. 7. 8. 5. 3. 4. 1. 2. lowing pages. lowing pages. your take exam.) to business hours It's time for your second exam. Locate a proctor area Form second exam. in your on the fol complete the Exam Request and your It's time for online at eLearning.uaf.edu and in the course received.online at eLearning.uaf.edu materials you own words understanding, be clearyour and show are imagineto you that explaining (To list. just do so - don't "describe" something, someone who of has no knowledge the subject.) Includea Study the information presented for this lesson until you feel that you have a good understanding. Write your answers your information the understanding. have a good Study Write feel that you presented you until lesson this for 122 BIOL F150: Introduction to marine biology Exam Request/Form 123 - - UAF eLearningUAF (Note: photocopying or taking notes from any examination paper or taking notes from is not permitted); examination any (Note: photocopying UAF eLearningUAF violation of process. the examination verification of instructional materials (if process; allowed during the examination any) return of all papers, including scratch and the completed Examination Cer questions, examination sheets, tificate to termination of confiscation of the examination, and immediate notification of materials, exam telephone ifby there is improper conduct on the part of evidence that there the student or any been a has examination session; examination identification of (or other verification, if photo I.D. the student by necessary); provision of area a quiet, well-lighted as free from noise and distraction and within supervisory as possible distance of the proctor; security of until it is opened in the student’s presence the sealed examination at the beginning of the • • • • • • Please sign the student’s Examination Request Form beforePlease sign the student’s Examination Request s/he returns it to our office and keep this Information referenceLetter for when administering the examination. advance by contacting UAF eLearning contacting UAF friends cannot be and/or advance Relatives by at 907-479-3444 or [email protected]. proctors. Proctor responsibilities include: stand their responsibilities and agree to ensure that integrity. When an individual is accepted as a proctor, s/he When an individual is accepted as a proctor, stand their responsibilitiesand agree to ensure that integrity. represents the University of process. the examination for and is accountable Alaska Fairbanks Proctors or an administratorcommunity college schoolat a public site be education officials must at a university, such other governmental persons or community officials, or if, or library, are unavailable, other people approved in You have been designated as a proctor Independent for Learning examinations. You The integrity of process the examination is fundamental to our program supervised because it provides the only thereforecheck are We of concerned that prospective proctors and capability. the student’s knowledge under INFORMATION LETTER TO BE GIVEN TO YOUR PROCTOR YOUR TO GIVEN BE TO LETTER INFORMATION 124 BIOL F150: Introduction to marine biology Exam Request/Form 125 Exam 2

http://elearning.uaf.edu Date: Phone: Phone Examination No:

UY4

Fairbanks, AK 99775-6700 Fairbanks, University of Alaska Fairbanks Section No: 907-479-3444 fax: 907-479-3443 UAF eLearningUAF & Distance Education

2175 University 756700 Avenue South, Suite 200 , PO Box EXAMINATION REQUEST FORM REQUEST EXAMINATION

Introduction to Marine Biology BIOL F150

The exam will be mailed directly to the proctor, so maintain contact with that person. so maintain The will be mailed directly to the proctor, exam request submit an exam from eLearning may the UAF you website: Alternatively, Send this request form in a pre-paid provided with course envelope or to: materials Select an approved testing site and arrange individual an appointment to proctor with an eligible the examination. Ensure the proctor that has read the attached Letter of Information Form. Schedule this Request and has signed farthe exam enough in advance receipt to allow for of preceding all lessons of and the mailing the exam the test itself. If area live there in the Fairbanks you this form. need to complete is no office Come to our during regular business hours exam. to take your 4. 5. 3. 2. 1. Student's Email: Student's signature: Student's Address: Information Letter, and agree to observeInformation Letter, all the requirements of process. the examination Proctor’s signature: I agree before taking it. examination, this to complete all lessons covered by Proctor's Email: I have read and understand the duties and responsibilities of proctor an examination as described in the attached Proctor’s Name & Title: Proctor’s Organization: Mailing Address: Student Name: Course No: Course Title: PLEASE PRINT OR TYPE INSTRUCTIONS 126 BIOL F150: Introduction to marine biology Part three Putting It All Together: Marine Ecosystems 128 BIOL F150: Introduction to marine biology Lesson 12: Life at the Top: The Epipelagic Realm 129 - - Ma Course

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it the lesson)

9th Ed.). Written Assignment Reading Assignment Reading . Climate Change on ArcticClimate Change Peoples") Online Learning at the publisher's can be found Center: www.mhhe.com/ castrohuber9e Understand characteristics the physical of the epipelagic and environment, the challenges and opportunities present the organisms they for living there. Know the types of organisms in the epipelagic, found and their adaptations life in this environment. for Be familiar with some of the important trophic relationships among the vari ous members of epipelagic communities. the information of pages Read this lesson in the following for this Guide Complete the rine Biology Complete the eLearningto UAF & Distance Education. C&H: Chapter 15 (pp. 332-359) and reread "Eye on Science" p. 431 ("Effects 332-359) and reread of on Science" p. C&H: Chapter "Eye 15 (pp. More information, including interactive flashcards, quizzes, videoclips and links, 1. 2. 3. 1. 2. 3. Objectives of this Lesson of Objectives lesson the completing steps for assignment Reading Life At The Top: Top: The At Life Epipelagic The Realm beganAlthough the epipelagic to humans since they been accessible has time, we still Darwin’s and has been studied at least since sailing the seas, have much to learn about this realm. 12 130 BIOL F150: Introduction to marine biology N coplankton also include some coplankton also include too small tobe caught in standard thetiny of bers mem are They lifeonEarth. of forms the mostnumerous as all phytoplankton, and rank to constitute the majority of rococcus minute of numbers Recall (Lessons 5 and 6) that thisvast realm. about ied atleastsince Darwin’s tolearn time,westill have much sailing the seas,they andhas been scientifically began stud has been accessible to humans since Although the epipelagic humans. andmammals,by including birds land-dwelling into coastal marine environments, andis exploitedrents by cur It alsois atnight carried to feed). the epipelagic upinto venture oceancreatures ocean (andmany deep life in the deeper sinks down, supporting at the surface matter produced other environments as well. Organic notonlybut in atthesurface consumers ton supports ton mostly microscopic of by numbers vast is teeming with life, all supported sunlit epipelagic The complex and unique ecosystems. the environment to form other andwith with each interacting specifically adapted, are organisms to which ments hasdistinct physical characteristics these environ tidal and subtidal zones andestuaries. of Each coastalenvironments,the inter we will explore 15, including ocean and,in and Lessons 14 lesson, we move tothe deeper the ocean, in the of surface the life near In this lesson, weexplore munties they support. and theunique biotic com specificmarinehabitats to explore . Primary production Primary major groups of marine organisms, we are ready ready we are marineorganisms, of groups major the the marine environment, and surveyed istics of character ow that the general we havedescribed , were unknown until fairly recently.unknown until fairly were , Yet they appear cyanobacteria picoplankton Life At The Top: The Realm Epipelagic , Synechococcus by phytoplank theepipelagic epipelagic zone epipelagic archaea (C&HFig.Table15.2, 15.1), plankton nets , an entire , anentire and phytoplank . In the next Prochlo domain . The pi . The • ------

they play an extremely important role in the marinefoodweb. role they play important anextremely that by learning most scientists, organisms weare be living femtoplankton smaller than picoplankton, the ton even plank of another category necessitated the addition of This things) bycoplankton (and all other living to 100:1. 10:1 that discovered recently ago. decades until a few And we have unrecognized life of that conditions appear to be in a state of rapid change. change. rapid to bein a state of that conditions appear now is critically important and globalrelationships regional these of A better understanding ocean andthe atmosphere. the deeper between theepipelagic, interactions biological thecomplex physical about and We tolearn alsohavemuch clear. within these communities more become interactions andthe complex discovered life are of forms asmore evolve bysignificantly altered these discoveries, andcontinues to communities has been epipelagic of Our understanding larger when food web the epipelagic of the rest DOMenters from ergy en The predators. for larger as prey toosmall to serve are which bial loop by consuming these bacteria andarchaea, zoan the food web. larger, Somewhat but still microscopic, into consuming bringing DOM, it back the vital function of minuscule thepicoplankton, of foodweb. the epipelagic Other members of significant part loop is a very so the microbial production, primary all the epipelagic of as half accounts foras much loop the into through the marinefoodweb back cycled matter( dissolved organic release seawater) of million 10 in a drop (about rine viruses ma of Phytoplankton cells infected by numbers thesevast (C&HFigs.Weand 15.25). 15.4 that DOM now realize members of the of members heterotrophic net zooplankton . Although viruses are not considered to notconsidered are Although . viruses nanoplankton marine viruses DOM bacteria and archaea, perform perform bacteriaandarchaea, ) into the ocean, which isre ) intowhich theocean, consume the protozoans. completethe micro virioplankton outnumber the pi microbial microbial proto , or ------131 Lesson 12: Life at the Top: The Epipelagic Realm

------. The halo seasonal keeps the bloom ice-edge blooms ice-edge .) Arcticcod areim an stratification convergent evolution releases fresh water, in contrast in resalt to the releases freshwater, and setting the stage for a phytoplankton a phytoplankton for the stage and setting zooplankton to be present,most of then the energy from Ifbenthos. the for becoming fuel sinks, bloom ice-edge an the spring whenlater in the ice melts the water has warmed sup bloom ice-edge the zooplankton, then for up enough portssuch Zooplankton and as krill a rich pelagic web. food copepods graze and are the phytoplankton turn eaten in by larger invertebrates, seabirds fishes, and by jellyfish, like and bowheadincluding seals and mammals ringed whales. on seals and other whales,on seals and other seabirdschickseat the that and eggs of other seabirdsand na gulls) glaucous (e.g., tiveon fish, seabirds, peoples subsisting and whales. seals the melting the ArcticDuring spring and summer, ice pack brine rejectionfresh As freezing. during leased by water spreads ofcreates it fromice edges, a layer melting the less dense waterfloats that surface,the at producinga cline sea ice releasesmelting ice algae, which seeds the bloom. organic accumulated the All debris of sympagicre life is Ifthe bloom. leased as well, fueling the waternot disturbed is a strong verticalby event,mixing at the surface wherephytoplankton grow they quicklythis in nutrient-richsunlit, These environment. are both pelagicthe Arctic. for in life critical and benthic If too earlymelts ice the whenthere spring, the in sufficient is waterthe many but too cold for is photosynthesis for sunlight their blood, whichblood, their preventsdamage from freezing. (Some unrelated Antarcticproducefishes another AFGPs, similar ofexample portantcod, Green Atlantic source, largerfood (e.g., for fish Arcticand char),halibut land kittiwakes, seabirds gulls, (e.g., fulmars, murres spotted ringed, (e.g., seals and guillemots), belugaribbon)and whalesnarwhal).and and (e.g., They eventhe diet of supplement polar bears, and Arcticfoxes areas in populations, human local whereare they washed ashorelarge in numbersstorms, winter are by and they Thus, energy some fisheries. taken by transferred is from grazerswaterthe to ice the within life by column, ice eating Arcticalgaecod, whichbeing eaten by and themselves in turnfall prey trophicat higher to animals Additional levels. trophiclevels consumersnetting arefisheries added by of polar bearsfisheries), salmon Arcticcod (e.g., consum primarytheir ing whaleskiller seals), preypreying (ringed , ------) in ), a com brine AFGPs (or sympagic Boreogadus Boreogadus saida (more200 species than ( (unique) to sympagic (unique) com gammaridean amphipods Diatoms , whose physical characteristics , whosephysical we Arctic cod endemic formedprocess the during of , including tiny flatworms,, includingtiny nematodes, antifreezeglycoproteins sea ice are the major producers sympagic in webs. food . These are viruses, inhabited by ofa multitude brine channels ice meiofauna major food sourcemajor food of TheseArctic species in webs. fish key food marine oily small, are extremelyand have a widespread, abundant circumpolar Ranging farther distribution. north spe other fish any than carry they cies, grazing on algae, others preyingon grazers or on each oth Some species are er. some example for munities, whichgraze large in numbers underside on the of Arcticsea copepodsareother zooplankton ice. Amphipods, and the of several extend ice, the and algalmay filaments meters waterthe into Arctic,the In column. ice algae can contrib halfbetter than ute ofprimarymarine total the production. organismsMany thrive on the underside of the ice, some crustaceansand rotifers. fromhave been identified Arcticand other types ofsea ice) ice algae algae areto grow able the in fromInsulated air, cold the lowermost ice layers. Mats of algae grow on the underside ute rejection microbes flagellates), (bacteria, diatoms, and an assortment of global weather systems and climate)global weather systems ofis great concern. of communities Unique organisms, called livingsea ice and at the ice- within can be found munities, water interface. Sea ice is infiltrated a network by of min vironmentswhere climatechange progressed.most is Spe extreme adaptedthese life in environments for superbly cies are facedrapidly with changing and the futureconditions, of uncertain. is many changesaffectHow these will complex the interactionsand ecosystems (and communities sustain that productiveEarth.on environments They are the among also because ofknown least well difficulties the of and accessing workingare environments ice. Polar areasin dominated by extrememost among the Earth on and are to the subject most extreme They fluctuations. seasonal are also the en Life In And Around Sea Ice Sea Life In And Around extremely important but A specialized part of the epipelagic is environment arethese some of Lesson 3. studied in Surprisingly, the most 132 BIOL F150: Introduction to marine biology regions can be hugely productive, teeming with jellyfish, can behugely mol productive, regions for example in regions, the Bering Sea. These tinental shelf over shallow con and retreats the ice edge advances where important is particularly effect “trickle-down” bloom. This by anice-edge lifesupported pelagic the flourishing from matter that sediments down the organic also benefit from during the season. Benthic communities thousand kilometers than a may move iceedge, more which low the retreating efficiently.more andmammalsall fol fish, seabirds Foraging toforage allows consumers and time,which bothspace of in terms their production, Ice-edge blooms also concentrate associated with melting sea ice. nutrients and stratification edge blooms, the blooms donot benefit from open-water foodweb. tothe pelagic energy However,ferring unlike ice- consuming zooplankton quickly the phytoplankton and trans may bealater at all.There noseasonaliceformed fuel photosynthesis,if or sunlight to is sufficient in the spring, there before too early the seasonal ice retreats An ice-edge bloom does not occur if aswell as for whales, human fisheries.and humpback like sea lions, killer whales predators, the largest even for targets worthwhile fishes are fishes andlarger age in huge soboth for schools, fishes migrate Most pelagic their own kind). many on juveniles of fish species prey of nile salmon eat the smaller juvenile smelts (and the adults onjuvenile salmon, andjuve some adult smelts prey while quite complex. For example,adult salmon eat adult smelts, species, like salmon.Feedinglarger can be relationships smelts,and sandlance,aswell as thejuveniles of capelin cod, Arctic and belugas.fishes include Pelagic whals forage like seals,nar fishes, and whales to seabirds, larger prey fishes forage is amplified byfoodweb Arctic smallerfishes, called This than any othermammal. years) than 100 longerlive (more slowly and tiny may more zooplankton.They grow of bers num out vast and straining water ing huge mouthfuls of size by toattain able their tremendous engulf ist, they are any toex animal ever With the biggest mouth of perhaps m long). (upto4 any whale the longest baleenplatesof of to 720 460 mouth carrying bow-shaped an enormous this bulk is taken upbywith thehead, of than athird More tons. as 75–100 huge,weighing as much are Bowheads open-water bloomopen-water , which eat zooplankton and then become , which with large numbers of of numbers with large ------formation of ice at polynyas produces a surface layer of layer iceatpolynyas asurface of produces of formation repeated and soon.The temperatures, to freezing water then the iceis blown freezes, away, water exposing more as theexposed ice-making machines, become features coastsandislands. These side of the leeward away from blowing winds keep ice prevailing where ter in the ice pack enon associated with sea ice: anotherphenom Some benthic communities benefit from its annual retreat. andthe timing of the seasonal ice pack, of ultimately on the presence depends er). All this productivity freez in your thefish sticks localsupermarket of jor source today (a ma in the world single species fishery is the largest fishery Bering Sea The fisheries in the world. crative the most lu some of thic species (halibut,support pollock) fishes (salmon) and ben bothpelagic populations of Large andmarine mammals. fishes, seabirds luscs, crustaceans, sonal ice pack is gone, and usually un sonal theyice pack fast come ashore prey,and toaccesstheir primary ringed seals. When thesea on, andrest seaiceto travel but require cellent swimmers ex andhumans. are Polarlike polar bears bears predators, other from under the ice to escape can dive whales. They like killer molt,andavoidmarine predators the ice torest, need tohaul out on Seals and walruses tion by polarbears. the pups toweatherandpreda of the exposure es, reducing out Ringed seals carve lanugo with soft downy fur—the born most pagophilic seals are of almost two months forringed and Weddell young seals. The takes until weaning, unable toswim which cases the pupsare their pups on the ice, and in many tic. Most ice seals nurse Weddell, seals in the Antarc and leopard Ross, crabeater and sealsin the Arctic, and harp bearded hooded, ribbon, pack. “ while andfishes, and otherinvertebrates cod, krill, Arctic Antarctic asa just under the ice serves ronment Sea ice is a in the same placeyearpolynyas afteryear. that tend to form at large generated gions may on these currents, depend benthos in some re down tobenthic environments.The nutrients surface that carries sinks, setting up a current (through densesalty water very Pagophilic ice seals —which is white in some species, is like white ringed seals.—which critical habitat ” and walruses give birth on top of the ice on top of birth give ” and walruses (ice-loving) seals include ringed, (ice-loving)spotted, seals include lairs in snowdrifts or pressure ridg orpressure in snowdrifts for many organisms. The envi for The many organisms. polynas brine rejection nursery ground nursery , areas of openwa of , areas ) which ) which for ------Lesson 12: Life at the Top: The Epipelagic Realm 133 , ------, affecting the , also depend Ringed seals Ringed polar bearspolar also are imperiled by shrinking also are imperiled by trophiccascade have been crowding onto shore in walruses black guillemot black Some scientists fear the majority ofthat Some scientists the world’s polar bears, includingoffthose all coast of the be gone will Alaska, and Wildlife Fish 2008, or sooner (in U.S. the mid-century, by Service officiallyas threatened). them listed Arcticseabirds such as the must Antarcticthe Like they penguin, Adelie ice conditions. range within nest of the productive can feed so they ice edge chicks.and their themselves Longtermof studies black guil when werethey Chukchialong the out hauled Sea and Bering shores. In 2009, 131 event.died a in similar Walruses need platformice an thickto supportenough greattheir weight, waterover positioned needs to be and it that enough shallow can dive down to the bottom to feed. they The also ice edge carries great them freshover distances foraging grounds, retreatsas it northbecom ice is the Now, spring. the in and is recedinging thinner deeper over and deeper water. requirealso seals Ice thickis that ice persistsand enough stages ofcritical to complete the them for enough long birth and rearinging ofmolt. the annual and pups, and widespreadthe most abundant the marine mammal in northprey and the principal of on snowdrifts lairs.their for Snowdrifts have been melting whenthe pups are dependent still on these protecearlier, Longtermtive caves. of studies polar bears Bay Hudson in are revealingand reproductivecondition their that rates also are decreasing,the decreasing along with seasonal ice pack. at the ice edge, which ice edge, at the receding is now farther earlier. and Thesechanges are surelyproducing others ripple they as througha in community the predators, prey competitors and of the altered populations. Theregreatalso is concern Arcticfor sympagic communities, pelagicthe and supportedcommunities benthic and ice- by view in of blooms, edge the dramatic declinesArcticin sea Some changes arewe areice that witnessing. already being forming been thinning, sea ice has seen. As later and break ing up earlier, unprecedented numbersofto tens (up at earlier thousands), areas and in wherebeen seen have not they the year, in times beforenorthwest on the coast of areOn land, animals Alaska. to greatersubject travel must stress, and they greater dis tances to reach foraging areas.the summer of During 2007, several walruses young mostly thousand stampedes, died in ------, which Adelie penguins and , whichrequire their sea ice for Emperor Antarctic krill evolved to coincide with the proximity of evolvedproximity the with coincide to sourcefood their krill-dependentpredators already arethe Among declining. hardestare hit feed exclusivelyThealmost on krill. of location nest their oftiming and the colonies, ing breeding their cycles, have local implications for the organisms the for implications local living regions.these in partsIn of Antarctic the where popu declining, sea ice is oflations nursery grounds, greatly have decreased and arere being which salps, inhabitwarmer typically placed by waters. Some Sea ice is melting, as we learnedmelting, is Sea ice global 3. With Lesson in warminghappening as fast twice restpoles as on the at the ofthereplanet, the a dramaticbeen has decline ex the in and thicknesstent of sea ice, and the ice packfreezing is These changes have enormouslater and breakingup earlier. pack and in polynys, or in areas or in benthic for enough pack shallow polynys, and in feederswalruseslike and bearded to reachseals bottom the whendiving from the ice platform. by used ice also is Drifting migrationfor animals and transport new to feeding areas. Crowding is reduced and the environment is cleaner. The ice Crowdingreduced is cleaner. is the environment and waterthe shields below fromextremely harsh the weather in regions,these and pressure and snowdrifts ridges provide aboveshelter easy access to food, ice. Sea ice allows the whereespecially concentratedis it ofedge at the ice the Sea-ice environments offer Sea-ice environments advantagesother critical to the livinganimals there. Comparedareas to land available for out, the ice provideshauling a vastof amount space and a variety of different food, for competition less with habitats, fromand isolation most predators and other disturbances. ice provides access to food for the Inuit peoples inhabit Inuit the ice provides for access to food Arctic,coastal the ing whosurvivedhave of thousands for walruses,yearsseals, hunting by whalesand polar bears. ciated with sea ice, in leads or polynyas or at the edge ofedge at the or leads or polynyas sea ice, in ciated with the three All ice pack, where belugas and narwhalsalso winter. cetaceans are environment, adaptedthis life in for beautifully lackingdorsala ofa thick layer by insulated fin, and blubber capableof sea breakingthrough to breathe.ice the Finally, til the ice formsice the til again fall.the in They breedpack,ice the on as on as well sea ice dens on winter and femalesmake their ventureland. Arctic ice, preyingonto the out on ringed foxes remainsthe scavenging seal pups and of polar by killed seals bears. Thebowhead whale entireits spends closelylife asso 134 BIOL F150: Introduction to marine biology bance and loss of habitat, at any stage of their life history. atany habitat, of stage bance andloss of by pollution, distur may be harmed marine organisms cod and other and to pollution. Arctic water able to rough vulner particularly but makes the eggs spawning occurs, suited tothecalmunder-ice environment where tions are adapta These with an unusually thin membrane. eggs for cod, example,risks Arctic to marinelife. have buoyant disturbances and pollution that pose serious also create tion. Although economically these activites advantageous, and extrac shipping, exploration and oil and gas creased toin same time, the shrinking ice is opening these areas At the coastal villages torelocate. someArctic of habitants the in forced the coastlines and have already eroding fast are surges storm and dangerous. Greater difficult more water, in rougher andhas resulted shore making hunting away from farther themtotravel icealsoforces receding the of the subsistence livelihood animals threatens ice-dependent populations of Declining years. this population in recent of decrease shown a drastic Seahave Beaufort lemots nesting on anisland in the western Inuit people , as well. The , aswell. The ------and the regional and global implications of these changes. and globalimplications of and the regional changing, these ecosystems, howthey are of derstanding to ourun Continued study is crucial discovered. even are life unique tothese environments may they belost before is still in its infancy,regions of it is possible thatsomeforms ice-dominated doses. andin large quickly Since the study of it comes when to change to adapt for organisms difficult factor.at thepolesisalsoa change more It ismuch ity of rapid The effect. devastating food,can haveaparticularly cies an involved.Loss of are chain, in each links andfewer chains, becausefewer foodweb the whole on effect profound can haveamore one link in afood chain communities. andtropical Breaking totemperate compared lowspeciesdiversity,cause they tendtohaverelatively be vulnerable particularly High latitude communities are like Arctic cod, which many other species depend onfor many otherspeciesdepend which cod, like Arctic anchoring spe - - - - Lesson 12: Life at the Top: The Epipelagic Realm 135 biomixing and biological nutrient pump El Niño-Southern (ENSO) and La Oscillation Niña sympagic and pagophilic brine rejection and brine channels albedo critical habitat nursery ground endemic species trophic cascade anchoring species vertical migration redalgal spring blooms, HABs, tides, blooms, and eutrophication and open-water bloom bloom ice-edge nutrient-limited light-limited, limiting factor, and limiting nutrient and overturnstratification Antarctic and coastal upwelling, upwelling upwelling equatorial • • • • • • • • • • • • • • • • - - Tricho glossary terms glossary , and Synechococcus , rete mirabile rous zooplankton planktivorous nekton forage fishes protective coloration, countershading and self-shading DOM, detritus and marine snow microbial loop suspension feeders herbivorous/grazing zooplankton and carnivo Prochlorococcusdesmium and dragbuoyancy neuston and pleuston phytoplankton and zooplankton phytoplankton and megaplankton, macroplankton, mesoplankton, microplankton, nanoplankton, picoplankton and femtoplankton/virioplankton holoplankton and meroplankton epipelagic zone and photic zone zone neritic zone and oceanic • • • • • • • • • • • • • • • • your textbook. your Listed below areListed below terms some key Do a self-check this lesson. for understanding. for If you're unclear about the meaning of ofany these terms, re-read in this course the material and check textbook, guide and in your the glossary at the back of 136 BIOL F150: Introduction to marine biology online at eLearning.uaf.edu and in the course materialsyouonline ateLearning.uaf.edu received. andinthecourse a Include thesubject.) hasnoknowledge of someone who something,"describe" doso-don't just list. (To explaining that you to imagine are andshow your beclear understanding, own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 7. 6. 5. 4. 3. 2. 1. whale, polar bear,whale, ringed seal,salmon. amphipods, diatoms, Inuitpeople, jellyfish, gammaridean whale, killer guillemot, bowhead black cod,beluga, Arctic (but levels, notnecessarily including limitedto)thefollowing trophic identifyingat leastfive organisms: for theArctic, isshowninFig. toppedby 15.24 (p. web herring afood web 349).Usingtheseasamodel,draw a North inFigs. isdepicted food 10.14and10.15inyour web textbook epipelagic (pp.A typicalAntarctic 222and223), toavoidbeingseenandeaten. organisms epipelagic usedby adaptations coincidingwiththephoticzone,isplacetoseeandbeseen.Describethree epipelagic, The predators? epipelagic themostefficient someof Whoare carnivores. efficient makethemvery which by nekton, shared many adaptations epipelagic Describethree notplanktivorous. nektonare theepipelagic Most of nekton. planktivorous examples of several andgive organisms epipelagic of sizes. describethiscategory Briefly of nektoncomeinahuge range Planktivorous environment. found intheepipelagic protozoans typesof andnameseveral critical functionthey serve, food web. (Lesson5),play intheepipelagic animal-likeprotists acriticalrole microscopic Describethis Protozoans, zone. epipelagic theproductive out of ways thatplanktonmanage toavoidsinking Describethree thecurrent. against By definition,planktoncan'tswim Define, by thefollowing terms: comparingandcontrasting, j. i. h. g. f. e. d. c. b. a. cover sheet cover

, not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , red tidesandice-edge blooms red self-shading andcountershading mirabile loopandrete microbial DOM anddetritus andkrill copepods Trichodesmium holoplankton andmeroplankton picoplankton andnetplankton phytoplankton andzooplankton plankton andnekton Written Assignment for 12 Written Lesson Assignment and submit the written assignment to UAF eLearning & Distance Education according to directions found to directions & Distance Education according andsubmit the written assignment to UAF eLearning (Please make your completeandconcise) answers and Prochlorococcus • brief and clear , but, also complete . For example, if asked to asked For. example, if in yourin Lesson 13: The Netherworld: The Mesopelagic & Deep-Sea Realms 137 - . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). found at the publisher's Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Understand characteristics the physical of the diverse encountered environments below the epipelagic: the mesopelagic, deep-sea pelagic, and deep-sea benthic, including deep-sea hot springs and cold seeps. Be familiar with the types of organisms in mesopelagic and deep-sea found environ and behavioral and their anatomical, physiological ments, adaptations. Understand some of the trophic relationships within mesopelagic, deep-sea pelagic and deep-sea communities. benthic the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education. C&H: Chapter 16 (pp. 361–381) and "Deep-Water Coral Communities" (p. 321) Coral 361–381) and "Deep-Water Communities" (p. C&H: Chapter 16 (pp. More information, including interactive flashcards, quizzes, videoclips can be and links, 3. 1. 2. 3. 1. 2. Reading assignment Reading steps for completing the lesson the completing steps for Pelagic organismsPelagic in the midwaters found and deep sea are fascinating and otherworldly, bizarre-looking creatures with their harsh coping amazing adaptations for displaying environment. this Lesson of Objectives Netherworld: Netherworld: mesopelagicthe & deep-sea realms The The 13 138 BIOL F150: Introduction to marine biology B number of deep-sea species to beashigh million! as 10 deep-sea number of these findings, scientists some have estimatedthe from since thedescribed cies has 1980s. been Extrapolating spe deep-sea anew two weeks, on average, and every bottom by someexpeditions beenseen before, have never the thespecimens pulled up from of percent as90 much As on Earth. organisms of group maythe mostdiverse be benthos suggests that the deep-sea oceanfloor the deep astonishing. more What little on done sampling has been Biodiversity people have seen it. few although very on Earth, tebrate clinidens fish called the the ocean depths, atoothy little of life. One inhabitant of alsocontains an abundance environment on Earth, largest shown, however,the sea, anddeep that the mesopelagic in the nextwill explore twolessons. has Recent exploration the othershallow or and well-lit environments we epipelagic, less than in the is much here organisms of concentration the Indeed, pressure. andunderextreme cold anddark seem aforbidding placefor oceandepths life, isms). The organ these hugely abundant, diverse knowledge of of lack our of into together one unit is a reflection croorganisms mi inaccessible asgrouping environment (much relatively this immense and of ourignorance of reflection son isa in just that we can one les cover all these habitats fact The abyssopelagic the includes sea , where no sun at all penetrates. The deep sea realm searealm deep The nosun at all penetrates. , where elow the epipelagic zone lies the most vast andleast zonelies the mostvast elow the epipelagic (or known environment on Earth—the ), is the prime candidate for most abundant ver midwaters in the ocean depths is proving to be even to beeven is proving in the ocean depths bathyal benttooth bristlemouth , and the The Mesopelagic &Deep-SeaThe Mesopelagic Realms and ), a twilight world, and the twilight a ), world, hadal bathypelagic and hadopelagic The Netherworld: ( Cyclothone ac Cyclothone , mesopelagic abyssal zones. deep and • ------

he descended half a mile down in a tethered metal mile downin tethered a a half he descended WilliamIn the 1930s, Beebe. theAmerican naturalist, was environment life in its natural deep-sea to observe first The badlyotherwise, though mangledin manythe process. were up to almost 5 miles proved of depths from son and his crew up by specimens dredged Sir Thom thousands of sea. The in the deepest commonly assumedthatnolifecouldsurvive Until the HMS tethered and untethered robots. Still, less than one percent Still, robots. less than one percent and untethered tethered and and Chinese scientists, using both manned submersibles conducted by Canadian,Russian, Japanese American,French, exploration, deep-sea theof pace discoveries spurred These time in 1977. for the first observed were them, surrounding vents drothermal thePacific. later, of to thefloor made expeditions were Ridge the Study) during which Mid-Ocean Undersea Cyana aftpropeller. its large of overfloor,the sea driven by the angleandspeed controlling the submersible, deep-diving maneuverable first bathyscaphe designedbygeneration the In 1964, Auguste. Deep 7 miles down into the placeonEarth, deepest Navy Lieutenant Don Walsh their made historic trip tothe son Jacques andU.S.to the bathyscaphe.Piccard's In 1960, balloon attached agasoline-filled and ascendedby means of his spherical metal m by 4,000 to using weights attached of adepth reached sea.He into the deep descending untethered of his dream realized into the stratosphere, flight viouslythe first made hadpre who Piccard, theSwiss scientist In 1954, Auguste somany the "dancing"lights of sphere of the of was observed for the first time, in 1972. A few years years A few time, in 1972. for the first observed was , were used in , were less than 3 feet wide, and was astonished to see was and wide, lessthanfeet 3 Mariana Trench Challenger bathyscaphe , andthe astonishingly lush communities

Project FAMOUSProject Alvin expedition of the 1870s, it was the1870s,it was expedition of and the French submersible, submersible, andthe French bioluminescent , in, the , then dropped the weights , then dropped Trieste (French-American (French-American Mid-Atlantic , the second , Challenger Alvin creatures. creatures. bathy , was was , Hy - - - - Lesson 13: The Netherworld: The Mesopelagic & Deep-Sea Realms 139 - deep-sea gigantism chemosynthetic prokaryote meiofauna and macrofauna epifauna and infauna baitfall hydrothermaland white vent, smoker black smoker cold seep seamount deep-water corals and bioherm biodiversity • • • • • • • • • • old. deep-sea Unique such communties an these, holding as incredibleof wealth biodiversitywaiting to be discovered, are extent full the Uncovering activities. human threatened by to whichevolvedhas life darkthis in world mysterious de explorationscientific on but only pends not investigation, and resources.its also on our ability exploit we to manage how

, - seamounts glossary terms glossary which growcan to deep-watercoral bioherms Alvin also have been found at depth. have been found Lacking also zoo and and other destructive and other practices. Some forms of life hermaphrodites and male parasitism pheromones vertical migrator and non-migrator bioluminescence and photophores counterillumination eyes and bilobed eyes tubular oxygen minimum zone (OMZ) minimum oxygen (DSL) layer deep scattering main thermocline Trieste mesopelagic, abyssopelagic bathypelagic, and hadopelagic zones midwater and deep-sea dysphotic zone and aphotic zone • • • • • • • • • • • • • Listed below are terms some key this lesson. Do a self-check for understanding. for If you're unclear about the meaning of ofany these terms, re-read the material in this course and check textbook, guide and in your the glossary at the back of textbook. your more1,000 than moreand can be ft high 4,000 than years the vast majority ofwhich are be to a home unexplored, may of community unique organisms. Unique communities deep-waterthese environment, lightless their in xanthellae coralsform as known mounds ofsea floor the been observed. has many Unfortunately, deep-seaoverfishing, are environments damaged being by trawling before be lost can even they may environments to these unique be discovered. Each of the 30,000–50,000 140 BIOL F150: Introduction to marine biology online at eLearning.uaf.edu and in the course materials you received. materialsyouonline ateLearning.uaf.edu received. andinthecourse a Include thesubject.) hasnoknowledge of someone who something,"describe" doso-don't just list. (To explaining that you to imagine are andshow your beclear understanding, own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 8. 7. 6. 5. 4. 3. 2. 1. directly or indirectly supported by these primary producers. producers. by these primary supported orindirectly directly in these communities, theanimalsliving of Describeseveral producers? theirprimary scribe atleasttwo)? Whoare communitiesfound these flourishing (de are lifeinabundance. Where dosupport benthichabitats Some deep-sea benthos. thedeep-sea of biodiversity speciesalsolowinthisenvironment? Briefly discussthe islow. of there living Isthediversity organisms dance of sea, andthe abun in the deep life difficult makeall aspects of high coldandextremely pressure darkness, Perpetual undertheseconditions. that enable themtofindmatesandaccomplishreproduction organisms deep-sea of adaptations evolutionary populated.Describethree andsparsely dark seaisvast, deep The andmacrofauna)? sea benthos(discussbothmeiofauna thedeep- theusualfoods/feeding of habits are andwhat andotherbaitfalls. Whatisabaitfall, falls" "whale of form toafood thebottomstays bonanzainthe occasionally put,andthey treated are becausematerialthatreaches tage have aslightadvan floor sea.Benthosonthedeep-sea inthedeep living Very toorganisms littlefood isavailable tothesurface? back nutrients, waters, accumulatedindeeper andfor returning oxygen for replenishing responsible waters in deep-sea conditions?Whatglobaloceanphenomenonis havemadefor undertheseoxygen-starved living tions organisms twoadapta zone.Whatare inthemesopelagic inLesson3,occurs of oxygenThe welearned minimumzone,which it occur? andwhy does Whatisthis habit, inby engaged organisms. ahabit mostmesopelagic of day tonightasaresult from sonarsoundingsduringWorld War position II,changes of scatteringlayer), inthecourse DSL(deep discovered The andhowthey benefittheseorganisms. these characteristics Briefly describefour convergent evolution. of food—another example of of allow themtocopewiththisscarcity which anatomicalcharacteristics certain share fishes,distantly however related, allmidwater zone.Nearly epipelagic the thatsinkdownfrom production primary of ontheleftover crumbs dependent largely are organisms Mesopelagic cence. biolumines animals, intheiruseof anddeep-sea betweenmesopelagic differences nescence. Briefly discussthree theirownlight,aphenomenoncalledbiolumi for producing adaptations haveevolved these realms of inhabitants the sea.Many of downtothedeep nosunlightatallpenetrates whereas zoneisatwilightworld, mesopelagic The cover sheet cover , not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , Written Assignment for 13 Written Lesson Assignment and submit the written assignment to UAF eLearning & Distance Education according to directions found to directions & Distance Education according andsubmit the written assignment to UAF eLearning (Please make your completeandconcise) answers • brief and clear , but, also complete . For example, if asked to asked For. example, if in yourin ------141 Lesson 14: On The Shelf: Continential Shelf Communities . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). found at the publisher's Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Understand characteristics the physical of in the subtidal benthic environments zone, and the challenges and opportunities present organisms they for living in these environments. includingBe familiar coral with the subtidal benthos, reef and how they communities, are adapted life in these environments. for Understand some of the trophic relationships of coral reefssubtidal and other benthic communities. the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education. C&H: Chapters 287-305 and 307-330) 13 and 14 (pp. More information, including interactive flashcards, quizzes, videoclips can be and links, 1. 2. 3. 1. 2. 3. Reading assignment Reading steps for completing the lesson the completing steps for The subtidal zone is home to a variety of communities living on the continental shelf's surface, or muddy as well as kelp forests sandy and seagrass unvegetated, mostly beds, and most of the world's coral reefs. this Lesson of Objectives Continental Continental shelf communities on the shelf: shelf: the on 14 142 BIOL F150: Introduction to marine biology B than those lying beyond the shelf break, which westudied which break, than those lying beyond theshelf the world's as well as on the shelf's mostly unvegetated,sandy ormuddy surface, communities living of subtidal variety zone ishometoa The in the following and estuaries, lesson. will covered be zone in the intertidal associated with the shoreline, ritic pelagic about this lesson. We learned thesubject of tide line, are zone) in Lesson 12. Communities more closely closely zone) in Communities Lesson 12. more on the shelf:Continental shelfcommunities enthic ment over the littoral life in the water column over the shelf (the column overthe shelf lifein thewater kelp forests coral reefs coral communities in the ) zone, the part of the marine environ of ) zone, the part . The subtidal is a much smallerzone subtidalmuch isa The . continental shelf and seagrass beds seagrass subtidal belowthe low and most of and most of (or sub ne - - - • makes it especially vulnerable to exploitation makes byit especially vulnerable humans, accessible.generally has become Its proximity to land also scuba technology, the subtidal environment the advent of Since rainforests. tropical of the richness systems, rivaling all marine eco and complex of the most diverse are here found reefs coral The resources. oil andmineral and major fishing grounds, the world's however, containing most of the ocean, of lessons. part It is the richest in the previous uncertain future, as we will explore further in Lesson 16. further as wewill explore future, uncertain an and face theseactivities of as a result damaged been already reefs—have communities—in particular,coral shelf major fisheries and many continental theworld's Most of human activities. and topollution from and otherdamage - Lesson 14: On The Shelf: Continential Shelf Communities 143 mesenterial filaments and sweeper tentacles mesenterial filaments and sweeper planula zooxanthellae bleaching bioerosion eutrophication ocean acidification ENSO/El Niño recruitment competition and competitive exclusion facultative symbiont obligate symbiont and trophic cascade anthozoans and hydrozoans hermatypic and ahermatypic scleractinians and lagoon atoll barrier reef, fringing reef, cay coral key/sand pinnacle, knoll, reef, patch and algal ridge reef reef flat, crest, reef back-reef slope, slope and fore-reef slope and groovebuttress/spur formation • • • • • • • • • • • • • • • • • • • glossary terms glossary holdfast, frond stipe, and pneumatocyst sporophyte and gametophyte encrusting algae, coralline red algae, and calcareous green algae epiphyte forest kelp bed and kelp understory and algal turf canopy, animals deposit feeder and suspension feeder passive suspension feeder and filter feeder bioturbator substrate sediments and biogenous lithogenous benthos demersal epifauna, infauna and meiofauna/interstitial continental shelf and shelf break tidal currents and turbulence stratification sedimentation subtidal/sublittoral zone low tide line • • • • • • • • • • • • • • • • • • • • your textbook. your Listed below areListed below terms some key Do a self-check this lesson. for understanding. for If you're unclear about the meaning of ofany these terms, re-read in this course the material and check textbook, guide and in your the glossary at the back of 144 BIOL F150: Introduction to marine biology someone who has no knowledge of thesubject.) hasnoknowledge of someone who something,"describe" doso-don't just list. (To explaining that you to imagine are andshow your beclear understanding, own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 9. 8. 7. 6. 5. 4. 3. 2. 1. been negatively affected. been negatively ecosystemhas reef thecoral howtheirenvironment hasbeendisturbedandthe healthof discuss oneexample of and corals, reef tropical of physical requirements risk.Describethree atgreat reefs putcoral environment, which todisturbancesintheir sensitive specificphysical very andwell-being, conditionsfor theirgrowth andare very require corals Reef reefs. coral of andbuilders designers thechief are corals help, reef Although they havelotsof thesehighly communities. competitive of to besuccessfulmembers other(for spaceorfood), competewitheach other, andtwoways thatthey assisteach organisms cific ways inwhich teemingwithlife.Describetwospe are reefs coral allmarineecosystems, tropical andmostcomplex of richest The critical contributions tothisprocess. make corals, otherthanreef organisms, Describehowthree isa"teameffort." reef andsustainingacoral Growing themselves. way ability? ismostcriticalfor Which theirreef-building feed ways. describefour Briefly ways thattheseorganisms of obtainnourishmentinavariety Reef-building corals andkelps. killerwhales involving seaotters, seaurchins, Alaska, theAleutianIslandsof around cascadethathasbeen studiedinkelpforests Describethetrophic effect. community, theentire canripplethroughout these communitiesisdisturbed,theeffects cascade" causinga"trophic in feedingrelationships kelpcommunities. complex, highly Whenonesetof productive of conspicuous members themost are They andsubpolarregions. temperate coastsof alongrocky waters, incoldnutrient-rich Kelps grow with thesechallenges. tocontend thatseaweedshaveevolved adaptations and lightconditions, andcompetition.Describethree grazing, temperature action,variable subtidalcommunities, subjectedtowave inhard-bottom they are producers primary Major toattach. onwhich substrate ahard andrequire roots true seaweedslack Although plant-likeinappearance, orvisitors. bedsbenefittheirinhabitants ways thatseagrass three inmany other ways. bedsprovide Describe seagrass onseagrasses, feeddirectly organisms beds. Whilefew grass sea thick they canform soft-bottomsubtidalenvironments, where intothesandormudof grow roots plants. Their orflowering theangiosperms, of marinerepresentatives theonly truly inLesson6,are aswelearned Seagrasses, detritus. makeuseof andoneinfauna, oneepifauna these organisms, unvegetatedenvironments soft-bottom in the subtidal zone dine on detritus. Describehow two of of Most inhabitants environment,contributes explaining toitshighproductivity. howeach continental shelf the of physical characteristics theocean.Describethree of part sea,but itistherichest anddeep mesopelagic totheepipelgic, themarineenvironment, compared of smallpart accountsfor arelatively continentalshelf The , not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , Written Assignment for 14Written Lesson Assignment (Please make your completeandconcise) answers • brief and clear , but, also complete . For example, if asked to asked For. example, if in yourin - - Lesson 15: Between Two Worlds: The Intertidal Zone & Estuaries 145 . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). found at the publisher's Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Understand characteristics the physical of intertidal and estuarine environments, and the challenges and opportunities present the organisms they for living there. Know the types of organisms in the intertidal found and their zone and in estuaries, adaptations life in these environments. for Be familiar with some of the important trophic relationships within intertidal and estuarine communities. the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education. C&H: Chapters 11 and 12 (pp. 244-266 and pp. 268-285) C&H: Chapters 244-266 and pp. 11 and 12 (pp. More information, including interactive flashcards, quizzes, videoclips can be and links, 1. 2. 3. 1. 2. 3. Reading assignment Reading steps for completing the lesson the completing steps for In this lesson, we explore varying habitats with widely because of conditions, physical their location at the intersection of two very different environments. this Lesson of Objectives worlds: worlds: intertidal The & estuaries zone between two two between 15 146 BIOL F150: Introduction to marine biology I these coastal extent of andrising sea levels.The storms sion, drought, species, in addition donebyto the damage invasive ero mining, logging, pollution and farming, ming, construction, dumping, dam been subjected to dredging, draining, most severely,by affected have human activities.They themarineenvironments most directly, are These and lesson, and we studied in the previous which flats include Estuarine habitats rivers. of at theoutflow the landmeetssea, conditions able where ronment. forcoping with adaptations this dynamicextremely envi of exposed tothe air. awide variety have evolved Its inhabitants and in saltwater immersed marine environment, isalternately zone tertidal tersection of two very different environments. The environments.The different twovery of tersection their location at the in physical conditions, becauseof with widely habitats varying n this lesson, we explore and Estuaries mangrove forests mangrove wetlands , the most accessible and familiar part of the of , the mostaccessible part andfamiliar also are exposed tounique and vari alsoare the intertidal Zone &estuaries has been drastically reduced and reduced has been drastically , aswell as between two worlds: salt marshes salt seagrass beds seagrass oyster reefs. , mud • in ------, and warming the nights, and as the Earth's "kidneys," filter the nights, and as the Earth's and warming cooling the days asairconditioners, cies. also serve They Coastal wetlands provide acoast. within miles of 60 concentrated are whom of than half our own species, damaging more nities, weare By destroying theseenvironments and commu they are. how important is still declining. We belatedly learning are and, oncelost, it may not be possible to duplicate them. amongthe most complex ecosystems on Earth are These difficult. wetlands, very but itisproving and restore repair now underway to are flooding. Manyand reducing efforts andactlikea giant sponge, surges soaking up excess water our coastal communities. andstorm waves Estuaries buffer to vital thattheyprotection provide clear it has alsobecome tsunami, Japan Indiantsunami Ocean and 2011 as the2004 tsunamis such Sandyand devastating in 2012, Superstorm and in 2005 Katrina like Hurricane violent storms of wake them.Inthe through that runs the water ing andcleansing critical habitats for many spe - - - Lesson 15: Between Two Worlds: The Intertidal Zone & Estuaries 147 - - eutrophication islands tectonic estuary and fjord (glacier-carved estuary) mangrove forest (mangal), tidal/salt marsh, reef and oyster mudflat, negative estuary salt wedge tidal current and tidal bore outwelling brackish water euryhaline and stenohaline osmoconformer osmoregulator and active transport and osmosis anadromous and catadromous pneumatophore invasive species biological clock transplantation, removal and cagingexperi ment ecological community succession and climax predator keystone competitive and resource exclusion partition ing coastal plain estuary (drowned river valley) wetlands sand bars and barrier estuary, bar-built • • • • • • • • • • • • • • • • • • • • • • - glossary terms glossary anoxic hemoglobin feeder sessile holdfast, threads and byssal pneumatocyst, interstitial water rocky bottom and soft bottom epifauna and infauna meiofauna/interstital and bioturba animals tors suspension feeder and filter deposit feeder, vertical zonation upper intertidal, middle intertidal and lower intertidal substrate wave shock tide pool immersion and emersion desiccation dispersal low tide, high tide, spring tide and neap tide high tide, low tide, diurnal semidiurnal tide, tide tide and mixed headland and bay wave refraction intertidal/littoral zone active margin and passive margin • • • • • • • • • • • • • • • • • • • • • • • your textbook. your Listed below areListed below terms some key Do a self-check this lesson. for understanding. for If you're unclear about the meaning of ofany these terms, re-read in this course the material and check textbook, guide and in your the glossary at the back of 148 BIOL F150: Introduction to marine biology online at eLearning.uaf.edu and in the course materialsyouonline ateLearning.uaf.edu received. andinthecourse a Include thesubject.) hasnoknowledge of someone who something,"describe" doso-don't just list. (To explaining that you to imagine are andshow your beclear understanding, own words for this lesson until you presented that you feel Write Studygood havea understanding. the information your answers 9. 8. 7. 6. 5. 4. 3. 2. 1. man activities which have contributed to the destruction of these vitally important ecosytsems. thesevitally important of have contributed tothedestruction which man activities estuarine environments. Describetwohu along ourcoasts. Unfortunately, of percentage we havedestroyed alarge storms outflow, tousbecausethey filterandcleanse of theeffects andbuffer important extremely Estuaries are estuarine environments. use of andhowthey make thesevisitors on adaily Describetwoof or seasonalbasis, oratsomepointintheirlifecycle. thisbounty, of partake organisms high.Many nonresident estuariesisvery viduals, however, of andtheproductivity indi of by numbers large abundantly represented speciesare species. Those few hometorelatively Estuaries are outside thecommunity, thatinyour alsoinclude discussion.) matterissuppliedindirectly, organic by producers (If mudflats, andsaltmarshes. intertidal, environments: rocky and estuarine lifeinthefollowing intertidal thatsupport producers community. themostimportant Describe someof a of orindirectly, directly materialsneededby allothermembers andraw producers, theenergy provide Primary conditioninestuarineenvironments. this widely varying for copingwith organisms by different adapted three most widely inestuarineenvironments? Describethestrategies thelandmeetssea.Whatphysical fluctuates factor where interactions subjectedtoextreme Estuaries alsoare for typically lifeinthatzone. isadapted found there one organism zones, describehow briefly vertical the three zonation. For of each display habitats adistinct vertical intertidal Rocky orestuarinecommunities. soft-bottom intertidal in infauna soft-bottomhabitats. Describehowthisisaccomplishedby typesof three of intothesubstrate burrow to for organisms andsoft-bottom coastalenvironments istheopportunity betweenrocky One obvious difference shock. copewithwave two organisms Describehow intertidal. lifeintherocky of isanotherchallenge Water tidesandcurrents movement duetowaves, asolution. have evolved tidal organisms howinter twoexamples theseproblems, of give of and,for each causedby theproblems emersion Describe twoof asthey dealwithauniquechallenge: emersion. organisms intertidal shore rocky At lowtide,wecaneasily observe itsaccessibility. isthesmallestzonewithinmarineenvironment, but itisthebestknownbecauseof intertidal The Terms" for thislesson.) inthe"Glossary included puzzle are inthecrossword (All theanswers withthisassignment. theanswers) puzzleonthefollowing page, andsubmitit(oralistof Complete thecrossword cover sheet cover , not those of the textbook or Course Guide.Be the textbookCourse or not thoseof , Written Assignment for 15 Written Lesson Assignment and submit the written assignment to UAF eLearning & Distance Education according to directions found to directions & Distance Education according andsubmit the written assignment to UAF eLearning (Please make your completeandconcise) answers • brief and clear , but, also complete . For example, if asked to asked For. example, if in yourin - - - Lesson 15: Between Two Worlds: The Intertidal Zone & Estuaries 149 5 1 tolerant of a wide range of salinities extensions of mangrove roots, growing up into the air final stage of ecological succession (2 words, no space) carnivore that makes a disproportionate impact on the community (2 words, no space) devoid of oxygen dense mangrove forests collective term for marshes and other coastal environments, partially or completely flooded at high tide 4 7 1 2 3 4 5 6 8 DOWN 12 11 6 10 3 (Tear out and submit for grading) out and (Tear 2 Lesson 15 Crossword Crossword 15 Lesson crossword puzzle for Lesson 15 for puzzle crossword 8 9 www.CrosswordWeaver.com "high and dry" in the intertidal (opposite of "immersion") pattern of banding in rocky intertidal communities (2 words, no space) impact from water crashing on shore, in the intertidal zone (2 words, no space) process by which surplus detritus is carried from estuaries out into the open ocean water of intermediate salinity 7 9 ACROSS 10 11 12 150 BIOL F150: Introduction to marine biology 151 Lesson 16: The Human Factor . Marine Course Guide

(below) in C&H (Castro & Huber, (below) in C&H (Castro & Huber, (last page of (last page and submit it to UAF the lesson)

Reading Assignment Reading Written Assignment 9th Ed.). Changing Planet" (pp. 231-242) Changing Planet" (pp. Online Learning at the publisher's found Center: www.mhhe.com/castrohuber9e Be familiar with the resources of humans. are the sea and how they used by some ofRecognize the scientific, in economic, social and political aspects involved managing marine resources. Understand the impacts of and marine life, and humans on marine environments efforts to conserve and restore them. Grasp of the global significance of the extent ecosystems, and marine environments their destruction, and the importance of their conservation. the information of pages Read this lesson in the following for this Complete the Biology Complete the eLearning & Distance Education. C&H: Chapter 17 (pp. 383–404), Chapter 18 (pp. 406–423) and "Special Report: 383–404), Chapter 18 (pp. C&H: Chapter Our 17 (pp. More information, including interactive flashcards, quizzes, videoclips can be and links, 1. 2. 3. 4. 1. 2. 3. Reading assignment Reading steps for completing the lesson the completing steps for We are beginning only to recognizeWe the impacts of human activities on the world ocean, just as we are beginning only to explore this vast realm and to understand its significance in the global ecosystem that is Earth's biosphere. this Lesson of Objectives factor factor the humanthe 16 152 BIOL F150: Introduction to marine biology T with one species can have far-reaching effects, such asthe such effects, with one speciescanhavefar-reaching intricately interwoven, andthat interfering community are a of We among the members haveseen that relationships (Lesson 12). bycommunities threatened globalwarming (Lessonand sympagic 13) communities subjected to trawling benthic as deep-sea such being discovered, before even disappearing, some life are of less conspicuous forms of exterminated auk andSteller's sea cow,like the great been have already of Some well-known12). species havebeenpushed to the brink sea-ice conditions (Lessons and 11 deteriorating including habitat, over-hunting,mammals from pollution and loss of and the (Lesson 10); andindiscriminate fishing practices habitat, of pollution, destruction oil spills and other typesof of result asa andshorebirds seabirds over-hunting, andtheloss of sea snakes due to and some seaturtles all species of of status endangered or the threatened (Lesson 9); tofish populations damage (Lessons and15); from 14 reefs and coral forests beds, mangrove seagrass of destruction ouractivities: of as aresult by marineorganisms suffered consequences the disastrous We havealso noted someof (Lesson 15). andotherhuman activities al development coast of estuarine environments, asaresult important of (Lesson and the 14); destruction and globalwarming levels sediment pollution, increased including factors, of variety a due to habitats tocontinentaldamage sons and 12); shelf 3 (Les globalwarming of seaiceasaresult of the retreat extinction rine environments: the break-up of iceshelves and of rine environments: thebreak-up on ma human activities of to the effects references numerous have been there this course, hroughout threatened , such as some of the great whales. Others, whales. Others, thegreat as someof , such (Lessons 10 and 11). Far greater numbers numbers Farand 11). (Lessons 10 greater or endangered status of many marine status of The human factorThe human • - - - which in turn supplies oxygen to deep waters andnutri supplies oxygen waters in turn todeep which oceanconveyor, the great helps to drive and its formation for critical habitat manyice. Seaice provides organisms sea in the lossof resulting causeglobalwarming, gases greenhouse consequences. Increased motion a series of Disturbing son 14). the physical environment also setsin the Aleutian Islands (Les around in kelp forests foodweb cascade trophic the environment. The amount of nitrogen that we pump nitrogen amount of the environment. The gen the for of now responsible 70% ties are human activi theocean, of In additiontoalteringthepH all marine ecosystems. of the richest support which corals onforand molluscsfood, that werely and thereef-forming crustaceans many marine ecosystems, many species of of shells,diatoms andkrill, key including links in the foodchains with calcium carbonate organisms of to impedethe growth toxins. impactonmarinelifemay be Its most direct fects of nutrients and theef of alteringthe availability and rocks oceanchemistry, by the weathering of increasing aspects of at any time during other this Acidification period. changes than change of rate faster millenia, andatamuch of dreds for hun occur by the yearhigher than has 2100, occurred hydrogen ions may of in the concentration a 3-fold increase continues, this trend If in hydrogen ion concentration. crease in 30% toa corresponds overyears the last200 seawater ocean acidification is gases greenhouse increasing Another consequence of and melting seen at the poles (Lessons 3 and12). warming when Disturbances can escalateunexpectedlyglobal climate. aswell as profoundly influencing waters, ents to surface (forms of nitrogen useable by primary producers) in producers) useable by nitrogen primary of (forms positive feedback positive effect of removing a predator from the from a predator removing of effect (Lesson 3). The decreasing pH of of pH decreasing The (Lesson 3). occurs, as with the accelerated aswith the accelerated occurs, reactive nitro reactive ------Lesson 16: The Human Factor 153 ------, the U. S. Global Ocean Eco S. , the U. U.S. GLOBEC U.S. quences for all of all quences for Earth's includinginhabitants, ourselves. futureOur own depends on our understandingof these on our actions. and, systems morecomplex importantly, gratingthe Southernin Dynamics Ecosystems and Climate Ocean), parts of Program.Dynamics system we workthat vital is It to better understand whichsystems, high-latitude these and monitor are undergoingmost rapidthe changes and whichare criti important cally cycling nutrient productivityand for the in world ocean, and powerful influencesclimate.on global explorewe will lesson, this In furtherofimpacts the hu to and our attempts man activitiesmarine ecosystems, on manage these precious resources more carefully intel and are recognize beginningto fully only We con the ligently. sequences of we areas just activities, human begin only to explorening sea and to understand the significance its Earth'sis global ecosystem that the in biosphere. is It increasingly clear that we areofthe health imperiling the worldfar-reachinghave will this that ocean, and conse

- - - - - ) - - Fed (Inte COML , and the ICED in the ocean. ocean. the in Endangered Spe Endangered , the dead zones dead International Whaling Commission (Southernand Ocean GLOBEC) (United Nations Convention on the Law ofLaw the on Convention Nations (United Marine Mammal ProtectionAct , the UNCLOS (the Bering Sea Ecosystem Study) and in the south the and in Study) Sea Ecosystem Bering (the SO-GLOBEC BEST by the Sea), the 1982 1982 Sea), the the Law ofSea treaty the ratified yet not by effortglobal the is S., U. the regulateto ofuse world'sthe includingoceans, conservation of marine habitatsand life 1). Comprehensive(Lesson informationon marine eco polar regionsin systems being providedis norththe in by (Lessons 10 and 11). A key objective10 ofand 11). key A (Lessons biodiversity the as carriedsessment of the Census by out Marine Life ( projecttarget to is conservation efforts 5, C&H p. (Lesson 216). Attempts to lessen or to lessen reverseAttempts some of the damage caused activities human includedhave ofenactment the by the eral Migratory Act Bird Treaty cies Act formationof the into the air and ocean increasesthe air and ocean into of the incidence can rain, acid diseases, other cer and algalexplosive and which blooms coral inundate reefs, paralytic cause shell and createfish poisoning, 154 BIOL F150: Introduction to marine biology completing the exam. is noneedto complete theform. Cometo ouroffice duringbusinesshours to exam.) take your andcompletetheExamRequest following your theappendix. Form area After completingyour your textbook. of attheback theglossary guideandinyour textbook, andcheck thematerialinthiscourse re-read theseterms, any of themeaningof about unclear you're If for understanding. for thislesson.Doaself-check somekey terms Listed beloware • • • • • • • • • • • • • • • • • • • • • • polymetallic nodules polymetallic products marine natural genetic engineering seeding andsalmonfarming salmon ranching /mariculture Endangered SpeciesAct Treaty Bird Act Migratory Federal Commission tional Whaling ActandInterna Marine MammalProtection Oceans Commission 1996andPew Sustainable FisheriesActof ICED andBEST UNCLOS, EEZ,COML,U. S. GLOBEC,SO-GLOBEC, source re andnonrenewable resource renewable fish flour(FPC),mealandoil ghost fishing seine,and trawl gill net,longline, purse maximum yield sustainable andoverfishing optimaleffort fishing effort, fisheries andindustrial catch andby-catch stock/population species clupeoid fishesanddemersal finfish andshellfish impacts anthropogenic written assignment glossary terms for thislesson itwillbetimefor your in finalexam. Locateaproctor - - • • • • • • • • • • • • • • • • • • • waste andsolid waste heavy metals,radioactive treatment andsludge sewage desalination andsalinebrines sistent substance andper non-biodegradable biodegradable, (clathrates) methane hydrate (OTEC) tidal energy, conversion energy oceanthermal and artificial reef and artificial habitat restoration areas, marine protected extinction andextermination gered species species,threatened speciesand endan rare species species andinvasive native global distillation magnification biological ocean acidification house effect andgreen gas carbon emissions,greenhouse hypoxic/dead zone phication cascadeandeutro nitrogen nitrogen, reactive pollution thermal dispersants PCBs and chlorinated hydrocarbons hydrocarbons, (If you live(If you inthe Fairbanks area there Allow 4 hours for Allow 4hours - - - - Lesson 16: The Human Factor 155 - in your in your . For example, if example, . For asked to complete , but also , but clear and brief • (Please make your answers and concise) complete your make (Please and submit the written assignment to UAF eLearning to UAF assignment written the and submit according Education & Distance to directions found Written Assignment LessonWritten 16 for biodegradable and persistent biological magnification and global distillation and saline brines sludge eutrophication and ocean acidification nodules and marine naturalpolymetallic products invasive species and native species finfish and shellfish demersalfishes and clupeoid fishes trawls and purse seines optimal catch and by-catch renewable resourcesnonrenewable and resources EEZs and PCBs , ofnot those or Course textbook the Guide. Be

cover sheet i. j. k. l. e. f. g. h. a. b. c. d. Human activities greatly as a result affect not only marine ecosystems, of what we remove, also what but we add to the sea. Briefly discuss the effects of three types of marine pollutants. Of briefly discuss one that has made a par all the impacts humans make on marine life and marine environments, ticular impression (positive or negative) on you. Describe three factors, effort, other than fishing which have adversely affected fishery resources. (factors could phenomena) include or physical other human activities, Given the decline of fish stocks, and a growing human population, the practice of mariculture is gaining interest. Describe three problems potential associated with this industry. Describe the relationship between the size of a population ("stock") and the growth rate of the population. What range of what population size, and range of fishing effort, is most conducive or yield, to a maximum sustainable optimal catch? What has been the actual fishing effort of most of the world's fisheries in recent and what decades, impacts have fisheries made on fish stocks? Define, by comparing and contrasting, terms:the following Define, by 5. 6. 3. 4. 2. 1. Includea and in the course received.online at eLearning.uaf.edu materials you Study the information presented for this lesson until you feel that you have a good understanding. Write your answers your information the understanding. have a good Study Write feel that you presented you until lesson this for own words understanding, be clearyour and show are imagineto you that explaining (To list. just do so - don't "describe" something, someone who of has no knowledge the subject.) 156 BIOL F150: Introduction to marine biology Exam Request/Form 157 - - UAF eLearningUAF (Note: photocopying or taking notes from any examination paper or taking notes from is not permitted); examination any (Note: photocopying UAF eLearningUAF violation of process. the examination verification of instructional materials (if process; allowed during the examination any) return of all papers, including scratch and the completed Examination Cer questions, examination sheets, tificate to termination of confiscation of the examination, and immediate notification of materials, exam telephone ifby there is improper conduct on the part of evidence that there the student or any been a has examination session; examination identification of (or other verification, if photo I.D. the student by necessary); provision of area a quiet, well-lighted as free from noise and distraction and within supervisory as possible distance of the proctor; security of until it is opened in the student’s presence the sealed examination at the beginning of the • • • • • • Please sign the student’s Examination Request Form beforePlease sign the student’s Examination Request s/he returns it to our office and keep this Information referenceLetter for when administering the examination. advance by contacting UAF eLearning contacting UAF friends cannot be and/or advance Relatives by at 907-479-3444 or [email protected]. proctors. Proctor responsibilities include: stand their responsibilities and agree to ensure that integrity. When an individual is accepted as a proctor, s/he When an individual is accepted as a proctor, stand their responsibilitiesand agree to ensure that integrity. represents the University of process. the examination for and is accountable Alaska Fairbanks Proctors or an administratorcommunity college schoolat a public site be education officials must at a university, such other governmental persons or community officials, or if, or library, are unavailable, other people approved in You have been designated as a proctor Independent for Learning examinations. You The integrity of process the examination is fundamental to our program supervised because it provides the only thereforecheck are We of concerned that prospective proctors and capability. the student’s knowledge under INFORMATION LETTER TO BE GIVEN TO YOUR PROCTOR YOUR TO GIVEN BE TO LETTER INFORMATION 158 BIOL F150: Introduction to marine biology Exam Request/Form 159 Exam 3

http://elearning.uaf.edu Date: Phone: Phone Examination No:

UY4

Fairbanks, AK 99775-6700 Fairbanks, University of Alaska Fairbanks Section No: 907-479-3444 fax: 907-479-3443 UAF eLearningUAF & Distance Education

2175 University 756700 Avenue South, Suite 200 , PO Box EXAMINATION REQUEST FORM REQUEST EXAMINATION

Introduction to Marine Biology BIOL F150

The exam will be mailed directly to the proctor, so maintain contact with that person. so maintain The will be mailed directly to the proctor, exam request submit an exam from eLearning may the UAF you website: Alternatively, Select an approved testing site and arrange individual an appointment to proctor with an eligible the examination. Ensure the proctor that has read the attached Letter of Information Form. Schedule this Request and has signed farthe exam enough in advance receipt to allow for of preceding all lessons of and the mailing the exam the test itself. Send this request form in a pre-paid provided with course envelope or to: materials If area live there in the Fairbanks you this form. need to complete is no office Come to our during regular business hours exam. to take your 4. 5. 2. 3. 1. Student's signature: Student's Address: Student's Email: Proctor’s signature: I agree before taking it. examination, this to complete all lessons covered by Proctor's Email: I have read and understand the duties and responsibilities of proctor an examination as described in the attached and agree to observeInformation Letter, all the requirements of process. the examination Proctor’s Organization: Mailing Address: Student Name: Course No: Course Title: Proctor’s Name & Title: PLEASE PRINT OR TYPE INSTRUCTIONS

Appendix 161 appendix

Appendix 163 50% but The with just The fresh yet. plants relatively to creating most Earliest years ago. until a worms, fish plants mammals, Mammoths, on called land in went extinct. The Flowering years ago. sea weed and appear. on appeared and the oceans. solidifies. billion billion cockroaches Marsupials, primates, pigs, the land rivers and species and appear. trees in the largest mass are common period. and life the first hominids atmosphere is in billion billion Flowering appear. cone-bearing appear. plants, continents only starfish. mammals with reefs humans The fungi extinction go marine when of this the atmosphere. cools and the land. Rodents, of the the mass extinction sponges birds Early The species appear insects and large camels dominated to living Mammals appear. of corals, mollusks, No known from 1.4 like all of all and with sponge spread across Modern birds. butterflies the end Primitive giant many adapt Period ends and oceans. New species like 95% winged from today. the land. whales by and ammonia. are ferns, palm-like plants. appear. the land. extinct. to dominate Earth’s crust insects, complex the oceans. Amphibians Fish echinoderms like and when land with dinosaurs organisms Early appear. fish of new animal appeared. to accumulate in The Period ends is of the Neogene Earth are bacteria from 3.5 appear. crocodiles appear. cats, and many other plants the globe. and plants going oceans are full forms are algae grasses. to form. dominate on First bears and civilization life. continue dominate and and and plants the land fish, The land trees life scorpions, appear. sharks, bees and begins very different number years ago appear. Feathered humans) for the first time. forms. swamps on animals jaws appear. begun land multi-celled of the mammals spread across beginning beginning few survivors of the Permian pine large have No known moon populate dinosaurs Amphibians like extinction known of all Many Reptiles appear. Fish first trees the first forests. Spiders, bony water First appears. primitive A short time. First complex Oxygen Earliest life Earth is mostly methane and Major biological events Major biological The (early developed sabre-toothed 10,000 Rise cats, dogs, spread across Dinosaurs modern plants dinosaurs Dinosaurs small. common cycads, and The Geologic Time Scale Geologic – 200 251 299 359 417 443 488 543 – – – – – – – – – – – – – 543 2,500 2,500 3,800 3,800 4,570 354 417 443 488 145 200 251 299 of years ago Today 23 23 65 65 145 Millions Devonian Silurian Ordovician Cambrian Jurassic Triassic Permian Carboniferous Neogene Paleogene Cretaceous Period

Eon

Era Mesozoic Era Cenozoic Era Paleozoic

Eon Eon

Era

Eon Phanerozoic retrieved from http://www.mysciencebox.org/timelines — under creative — retrieved commons license. from http://www.mysciencebox.org/timelines Assembled from the International Commission on Stratigraphy 2004 report A Geologic Time Scale and from information on the Assembled from the International Commission on Stratigraphy 2004 report A Geologic Time is licensed under the Creative Univeristy of California Museum of Paleontology website (http://www.ucmp.org). This work Commons Attribution-NonCommercial License. To view a copy of this license, visit http://creativecommons.org/licenses/by- 94305, USA. nc/2.5/ or send a letter to Creative Commons, 559 Nathan Abbott Way, Stanford, California Archean Hadean Proterozoic Eon