Wednesday, February 1, 2012

February 1st, 2012 class

1. Introduction: Problems facing taxonomy and diversity.
2. Accelerating “taxonomy”: DNA barcoding.
3. Promoting taxonomy: Census of Marine Life.
4. Images from one CoML Project (Creefs).
5. Conclusions.
6. Test advice.

1. Introduction: Problems facing taxonomy and diversity.
Problems facing taxonomy 1
Too many species! Diversity confounds our best efforts to examine it.
Keep finding new species.
Extinction rates increasing.
Problems facing taxonomy 2
Not enough taxonomists.
Pay poor, work long.
Everyone says “important” but not considered essential.
Many groups have no active workers.
Potential solutions 1
Increasing technology and information available.
Global information systems.
Molecular experiment techniques.
Potential solutions 2
Increasing international research collaboration.
Growing awareness of biodiversity and importance.

2. Accelerating “taxonomy”: DNA barcoding.
What is “DNA barcoding”?
A DNA barcode is a short sequence, taken from standardized portions of the genome,used to identify species.
If a genome project is deep and narrow, DNA barcoding is broad and shallow.
Genome projectは深くて、狭いが、遺伝子バーコードは浅くて、広い。
Requirements of a DNA barcoding marker
A sequence/marker used to barcode should:
be easy to amplify
not possess paralogues
have conserved regions to design primers efficiently for a broad taxonomic sampling
be variable enough to distinguish species
but conserved enough within species
Choosing the correct DNA marker is critical.
Barcoding does not aim to solve phylogeny!
Reasons for DNA barcoding
1. Works with fragments.
2. Works with all stages of life: Can link male/females. Different stages of same organism. E.g. Amphipods (White & Reimer 2012)
3. Cryptic species detection. E.g. Astraptes
4. Reduces ambiguity (set DNA code).
5. Makes expertise go further.
6. Democratizes access to data. E.g. Barcode of Life project
7. Opens the way for handheld barcoders.
8. Finds new diversity.
9. Demonstrates value of museum collections. Sequencing of collections vital.
10. Speeds up discovery of new species.
Additional strong point
Does not need expert knowledge.
Weak points
1. DNA (specifically COI) does not always work for each group of organisms.
2. Handheld technology has not succeeded, despite many advances.
3. Different taxa have different DNA protocols, so standardization is difficult.
4. The “barcoding gap”
Barcoding implies that the level of DNA divergence between and within species is different.
But evolution not neat - hybrids, incomplete lineage sorting, etc.
This gap is not always present – so taxonomy comes back to “judgement”.

DNA barcoding proposed in 2003 as a “solution” to taxonomy.
Two large projects: Barcode of Life and Ocean Genome Legacy.
Encyclopedia of Life on the internet.
Common method of identification.
Gaining acceptance as a practical method to obtain much data.
But has not solved taxonomy, instead a new approach or “sub-field”.

For zoanthids (and corals), > 1 DNA marker is needed.
mt DNA evolves very slowly.
Still, better than no experts at all!

3. Promoting taxonomy: Census of Marine Life.

Scientific Framework
What has lived in the oceans?
What does live in the oceans?
What will live in the oceans?
The Census consisted of four major component programs organized around these questions.
1. Investigating the Past
Census researchers undertook the challenge of constructing the history of marine animal populations since human predation became important, roughly the last 500 years. This program component was called HMAP.
Teams of fisheries scientists, historians, economists and others conducted case studies in southern Africa, Australia, and approximately a dozen other regions.
Together, these case studies created the first reliable picture of life in the oceans before fishing.
The long historical records of marine populations help distinguish the contributions of natural fluctuations in the environment from the effects of human activities.
2. Assessing the Present
The largest component of the Census involved investigating what now lives in the world's oceans through 14 field projects.
Each sampled important kinds of biota in one of six realms of the global oceans using a range of technologies.
This included CReefs.
3. Forecasting the Future
To speak about what will live in the oceans required numerical modeling and simulation. This component program is the Future of Marine Animal Populations (FMAP).
Integrating data from many different sources and creating new statistical and analytical tools to predict marine populations and composition of ecosystems in the future
4. Living Legacy
Such a global initiative required a state-of-the-art data assimilation framework, and this is:Ocean Biogeographic Information System (OBIS).

2,700 scientists
80+ nations
540 expeditions
US$ 650 million
2,600+ scientific publications
6,000+ potential new species
30 million distribution records and counting

Example: CReefs:
Heron Island Trip
November 8th – December 1st, 2010
Census of Coral Reef Ecosystems (, part of Census of Marine Life (CoML –
Large international effort to understand biodiversity, use data for conservation.
Many researchers from different institutions, focused on different taxa, many “ignored”.
I focused on zoanthids (of course)!
Heron Island
Many seabirds nest here. Always noisy, and dangerous to walk without a hat!
Rails also live on the island.
The island is an important nesting site for green and loggerhead sea turtles.
Heron Island Research Station is run by the University of Queensland. Most of the station is very new.
There are excellent facilities for experiments, sample collection, and analyses, indoors and outside.
Scientists from all over the world, most based in Australia, but others from USA, Japan, Iceland, etc.
Most scientists had much field experience, and were good divers. Many also had boat licenses and first aid training.
There were also dive officers, who acted as boat captains, guides, diving assistants, etc.
Outreach included a professional photographer, and a professional blogger with stories and images posted every day.
A professional chef ensured everyone was well-fed. The food was amazing!
Every night at dinner, based on everyone’s ideas, weather conditions, and tides, the next day’s schedule was decided.
Boats were launched from the harbor, 3-10 people per boat. Often boats were out for over 7 hours.
Diving was always done under the buddy system, with very strict guidelines on diving protocol.
Special permits were obtained to allow specimen collection. Work underwater was intense and focused.
I was able to collect 270 specimens in 33 dives. Other groups collected up to >2000 specimens!
Samples were also collected on reef walks, and by snorkeling. Other groups used ARMS and the “carpet of death”.
Some groups found many new species, others new records for the southern GBR. This genus was found in the southern Pacific Ocean for the 1st time, likely a new species.
Another potential new species of zoanthid.
Many people also collected for other groups.
Back in the lab, data were collected, and specimens numbered. All data were given to the CReefs data manager as well.
All sites were assigned numbers, and all had GPS coordinates. 130 sites were visited in 3 weeks!
Samples will be shipped to institutions all over the world and analyzed further.
Of course, work was not 24 hours a day…
Sunset drinks were the one time of day when everyone would relax and take a break.
And the sunsets were amazing…

References cited:
1. CP Meyer, G Paulay. 2005. DNA barcoding: error rates based on comprehensive sampling. PLoS Biology 3(12) e422.
2. Consortium for Barcoding of Life homepage.
3. Census of Marine Life homepage.
4. OBIS homepage.

Wednesday, February 8th; 8:30-10:00.
10 questions, choose 7.
Open book, no cellphones, etc.
Must pass to pass class.
Arrive after 9 a.m. – out.
Advice: study!

December 21st, 2011 class

1. Introduction: What is a species?
2. Taxonomic species: Amphipods in the Ryukyus.
3. Biological species: Fish species complex!
4. OTU & barcodes: ARMS and estimating coral reef biodiversity.
5. Conclusions.
6. JCRS images.

1. Introduction: What is a species?
What is biodiversity? (Review)
Biodiversity = Number of taxa (species, genera).
OTUs = operational taxonomic units
Coral reefs are highly biodiverse, <0.2% of surface of Earth, >25% of marine species live there.
What is a species?
A species can be different, depending on who you talk to, or what meaning is implied.
Before we discuss species, we must be clear about what species concept we are discussing.
Three types are:
1. taxonomic species
A species with a scientific name.
Names are important for science & conservation.
Accuracy and authorship important.
Naming species takes time & money.
Few experts.

2. Biological species
A species proven to reproduce only with itself.
Reproductive isolation.
Species concept important.
Experiments take time & money.
Hybrids? Evolution is messy.

3. OTU/barcode species
A species defined by genetic differences.
Implies a threshold value.
Quick and easy.
Sequencing species takes time & money.
Limited by previous knowledge.
Not always “true”, barcoding gap is problematic.

It is estimated we have described 1~25% of the total marine species.
Description of all species – possible?
However, describing is important.
1. Unknown species - What kind of undescribed species exist?
We have no idea what awaits discovery.
2. “Hidden” species
Hidden species may be hidden due to their cryptic nature, or due to researchers not being able to distinguish them from other, similar species. Usually small.
Often, there are large groups of “species complexes” that are hard to make sense of.

C. Known but not described species
We know this is a zoanthid. DNA has shown us how it is related to other zoanthids (=very unique).
We also know it is undescribed, and only on Okinawa.
It is waiting to be described.

Part 2: Taxonomic species
Commensal amphipods in Okinawa
White & Reimer 2011-2012
Taxonomic species
Can be detected through observation of morphology or classic diagnostic characteristics. DNA?
Compare with previous taxonomic literature.
Only way to name a species.

Leucothoidae (139 sp.) – one family of amphipods
Anamixid clade (pronounced sexual dimorphism):
• 22 Anamixis Stebbing, 1897
• 4 Nepanamixis Thomas, 1997a
• 13 Paranamixis Schellenberg, 1938
Leucothoid clade
(minimal to moderate sexual dimorphism):
• 98 Leucothoe Leach, 1814a
• 2 Paraleucothoe Stebbing, 1899
Ecologically unique
• Endocommensal associates of sponges, ascidians, and bivalve mollusks
• Extended parental care
• Social system
• Potential eusociality

These are completely unexamined in Okinawa!
Contain two morphotypes, leucomorphs and anamorphs, previously thought to be different families of amphipods! DNA barcoding confirmed same species.
Materials and Methods
• Collect in-situ directly from host, take whole host, or coral rubble
• Traditional taxonomy
• Digital Inking
• Molecular DNA sequencing
25 new species from Okinawa in 1.5 years. More diverse in Okinawa than the GBR.

Part 3: Biological species
Fish species complex in the Pacific (Leray et al. 2010)
Biological species
Can be detected through observation of reproduction.
Also, reproductive experiments.
But these require extensive field work. Recently, molecular analyses usual, using phylogeny.
Background – Genus Dascyllus: Complex has 4 species:
1. D. trimaculatus (E. Africa to Central Pacific): 3 spots
2. D. albisella (Hawaii): white flanks
3. D. strasburgi (Marquesas Islands): gray
4. D. auripinnis (Line, Phoenix Islands): yellow fins
Each species has its own ecology and preferred habitats.
Materials and Methods
Specimens (n=563) from across the Indo-Pacific.
Microsatellite DNA data examined (data from previous research included).
Phylogenetic analyses and genotype assignment.
Previous research showed 5 clades, not 4 (Bernadi et al. 2003).
This research showed 7 (!) groups of Dascyllus, not all with clear morphology.
Appears ancient history of Indo-Pacific combined with recent evolutionary events drives speciation.
Hybridization appears to be occuring.
NOTE: unknown species are not yet valid species, or new species. They are undescribed species.

So…. Biological species
Results could lead to “finding” undescribed species or groups
This leads to…
Taxonomic species
Describing a species.
Only way to give a scientific name.
New species.

Part 4: OTU/barcode species
ARMS and crustaceans
Plaisance et al. 2011;
Brainard et al. 2010
Coral reef monitoring of other species asides from fish/corals.
Establish systematic and consistent metric to assess and monitor change.
ARMS – manmade deployment devices to look at crustaceans/molluscs, etc., simulate dead coral heads.
Easy to build, 200USD each.
Rugged, look like plastic boxes/crates.
Prototypes tested in Northwestern Hawai’i. Seemed to work well.
Deployed worldwide, and retrieved. Then processed, samples preserved, DNA sequencing (DNA barcoding).

So…. OTU/barcode species
Results could lead to “finding” undescribed species or groups
This leads to…
Biological species
Results could lead to “finding” undescribed species or groups
This leads to…
Taxonomic species
Describing a species.
Only way to give a scientific name.
New species.

References cited:
1. CP Meyer, G Paulay. 2005. DNA barcoding: error rates based on comprehensive sampling. PLoS Biology 3(12) e422.
2. KN White, JD Reimer – three papers (2011-2012). Commensal Leucothoidae in the Ryukyu Archipelago, Japan. ZooKeys (in press).
3. M Leray, R Beldade, SJ Holbrook, RJ Schmitt, S Planes, G Bernadi. 2010. Allopatric divergence and speciation in coral reef fish: the three spot dascyllus, Dascyllus trimaculatus, species complex. Evolution 64-5: 1218-1230.
4. L Plaisance, MJ Caley, R Brainard, N Knowlton. 2011. The diversity of coral reefs: what are we missing? PLoS One vol 6 issue 10 e25026.
5. R. Brainard et al. 2009. Autonomous Reef Monitoring Structures (ARMS): a tool for monitoring indices of biodiversity in the Pacific Islands. Pacific Science InterCongress, March 4, 2009.

Part 6: Japan Coral Reef Society 14th Annual Symposium
Images from the conference.

Tuesday, January 31, 2012


December 21st and February 1st class notes will appear later today.

January 25th, 2012 class

Presentations of applications by 15 groups. Overall, very well done; all groups passed.

Good job!

January 18th, 2012 class

With Kimura-sensei.

Human evolution

“genjin” left Africa 20-10 man years ago.
Then “kyujin” (Neanderthal), and “shinjin” (us) – Neandethals expanded range a bit, but we expanded a lot. We were able to adapt to many environments, changing genotypes and phenotypes, lifestyles too.

1. Evolution
2. Pop gen
3. Genome analyses – tough, but can learn about genome analyses, don’t have to learn all!

1. Evolution
Darwin and Lamarck intro. Could be good to distinguish between the two.
Important points: a. DNA to RNA to proteins. DNA info.
b. cellular structure
c. gametes.
Discusses allele selection and natural mutations. Most mutations disappear, but a few stay. Stay by either selection or random!

Neutral evolution: random mutation, not directly apparent on selection.
Natural selection: of course, as explained.

1.5-3 mya, very diverse time. P. bosei, P. robustus, P. ethiopicus, A. africanus, H. habilus, H. ludorufeo. H. ergaster.

Order: Use of tools, then into Europe, then into Asia, then use of fire.

Maybe different theories on human evolution. Diverse ancestors, or all one group, or reticulate evolution, or half and half. Humans are one group, with some small reticulate evolution.

H. erectus was alive until recently, and also Neanderthal was in Europe same time as H. sapiens. "Heidelberg man" alive until recent in Mongolia/Tibet.

2. Pop gen.

Humans have 23 chromosomes. Ome=all (zentai). Genome, proteome, etc. 22 normal and 1 X/Y!

Chromosome structure. Chromatid, telomere, centromere, etc. Also introduce DNA base pairs, AGCT. A and G purines, C and T pyrimidines. A and T bond, etc.

Types of DNA mutation.
1. Translocation – multimegabase to chromosome, or monosomy, trisomy. Change in ploidy.
2. Kilobase to megabase: Tandem duplication, deletion, inversion
3. 10s to few kilobases: Alu element insertion, minisatellite, element insertion.
4. Few base pairs: substitution, single indels, microsats. E.g. SNPs. Transitions more common than transversions (C to T, G to A).

Discusses DNA and inheritance from each parent, plus mutations. Polymorphism, and how some stay in populations, and some disappear. How this can also lead to new species. Can compare number of changes with molecular clock.

Y and mtDNA do not recombine.

Genetic drift and founder effect. Uses example of alleles in population (gene pool) and how it can happen.

Small sizes can have harsh genetic drift much more easily. N=20 always rapidly fixes, n=1000 not much but changes. Bottleneck and founder event.

For humans, mtDNA has lots of mutations. Y chromosome too.

Y follows paternal, mtDNA follows maternal. Therefore can follow wide range of evolution. Be careful making assumptions using only one side of DNA, missing lots of the story!! By doing the genome, we can look at the rest of your ancestors. mtDNA’s Eve is the same thing – be careful! Y Adam is the same.
MRCA=most recent common ancestor.

Thus, much DNA has incomplete lineage sorting. Adding more DNA markers and you can get complete lineage sorting.

2003 Human genome project finished.
Then, SNP project. HapMap.
Human Genome Gene Chip – 5man to 100 man SNPs (in one day). Next generation sequencer – soon a genome for 1000 USD.
Can envision reading all babies when born (!).

Genome-wide association study (GWAS); demography (pop movement etc/pop structure), natural selection, ancestral spp analyses.
a. GWAS: e.g. finding SNPs that link with human height. Weedon et al. 2008 etc. and related studies, now over 100 SNPs linked with height. Best to get all 100 SNP information, then calculate height.
b. Genome wide SNP, compare internal pop structure with distance from other populations. Infer pop history. E.g. Novembre et al. 2008 – mapping data fits with geographical history. Also migration in Polynesia; Tonga is mix of E Asia and PNG (Kimura et al. 2008; Price et al. 2008). Also examine admixed populations, understand how many generations have passed since admixing occurred.
c. Asia Pacific people came from E Asia. Several times of admixing/splitting.

Whole genome analyses:
Can use whole genome (n=2) of 1 person, can infer population size. With multiple people’s genomes, can infer sizes of populations in past. All infer serious bottleneck of 5000 people or so 50000 ybp. Do bottlenecks fit natural patterns? Usually to infer pop size, many samples, but can actually do this with whole genome! (Li et al. 2011).
Gronau et al. 201? – Bayesian inference from 6 genomes. All say humans evolved 50000 years ago. African populations split a long time ago.

Positive selection looks to fix must faster than neutral evolution (via selective sweep).

Population specific positive selection (selective sweep). Can look at human genotypes and phenotypes to examine this. Examples: Alcohol dehydrogenase, EDAR, etc.

Neanderthal genome – 1 to 4% in our genome!

January 11th, 2012 class

Conservation History on the Great Barrier Reef:

The Great Barrier Reef = GBR
Great Barrier Reef Marine Park

Why was rezoning of GBR necessary?
Representative Areas Program (RAP) (only part of solution)
Phase 1 and 2
Final zoning plan
Implementation phase
Other actions
Reef Water Quality Plan
Reducing fishing and policing

The Great Barrier Reef = GBR
345,000 km2
> 2000 km long
2900 separate reefs
> 900 islands

Formation of the Park
Late 1960’s – early 1970’s—much agitation for a park, reinforced by plans to mine Ellison Reef (off Innisfail)
Politicians promised that the GBR should be protected as a Park
Park established in 1975, under Great Barrier Reef Marine Park Act (Federal Parliament Act)
Park boundaries are non-negotiable, can only be changed by Act of Parliament
No mining within the Park
Development & implementation of zoning plans is a Federal responsibility
Day to day management is the responsibility of Queensland Parks & Wildlife Service
Zoning Plans
First areas to be zoned Capricorn and Bunker, finished in 1977
Subsequently the other regions were zoned
Zoning plans reviewed at regular intervals, with public participation, and plans changed over time and even the type of zones changed

Based in Townsville
Responsible to Minister for Science
Issues permits and licences, including those for scientific research
GBR declared World Heritage Area in 1981— such listing requires regular report card to ensure the reef is being maintained

During the 1990’s
Increasing use of the reef by tourists
Increased scientific knowledge of the reef
Increasing awareness of the connectivity of reefs (mass spawning)
Increasing evidence of decline of some habitats, especially inshore
The Great Barrier Reef Is ‘Under Pressure’

Downstream effects of land use (water quality issues)
Coral bleaching
Coastal developments
Increasing fishing effort and impacts
Shipping & pollution incidents
Increasing tourism and recreation
Trends in Regional Biodiversity Are Negative
Fishing effort increasing substantially in intensity & spatial extent (coral trout fishery—effort x2 since 1995; shark catch x5 since 1991)
Turtles–all 6 species threatened; 2 are endangered (Loggerhead and Olive Ridley)
Dugong population south of Cooktown has declined >90% since mid-1980’s
Humpbacks listed as vulnerable; other cetaceans (Irrawaddy & Indo-Pacific hump-backed dolphins) listed as rare
Trends for most species unknown
GBR Is Not Isolated From World Trends
10% of world’s reefs destroyed or severely degraded
58% of world’s reefs potentially threatened
70% reefs already degraded in Indonesia & Philippines
On current trends 70% of the world’s reefs will have gone in 40 years
Minimising the
Downstream effects of land use ==> Reef Water Quality Action Plan (results not immediate)
Coastal developments ==> Aquaculture Regs; GBRMP permit requirements
Increasing fishing effort and impacts ==> Queensland FS fisheries management plans (ECTMP, Reef Line)
Minimising the
Shipping & pollution incidents ==> Australia Marine Shipping Authority shipping review, compulsory pilotage, mandatory reporting, etc
Increasing tourism and recreation ==> PoMs; new tourism framework
Threatened species ==> new policies; species recovery plans; seasonal closures, RAP
Protecting biodiversity ==> RAP
Why was rezoning of the GBR necessary?

Queenslanders depend on the GBR
Important for economy—tourism, commercial fishing, recreational fishing, shipping
Important for Traditional Owners—connection with Sea Country
Important for communities—relaxation, lifestyles
>90% Australians (including Queenslanders) wanted more no-take zones
Important for building knowledge—education, research
Better protection = insurance for all these values

Connectivity in the GBR

An overview of RAP
Representative examples of the entire diversity of habitats protected
RAP reviewed the existing zoning of the Marine Park
RAP attempted to minimise negative impacts for users and stakeholders while aiming to achieve protection of biodiversity
RAP has meant an increase in Green Zones to protect biodiversity
RAP is a crucial part of the solution to a complex problem

Other Issues Addressed During Rezoning
Some current zoning plans had been in existence for 16 years
Ensured consistent zone names and zone provisions throughout GBR
Coastal areas zoned for first time
Clearer delineation of zone boundaries (GPS co-ordinates)

Developing the Zoning Plan
The Zoning Plan was developed using environmental, economic, and social information
Clear Principles on how to use the environmental and social information were followed
These principles were set out in the first round of community participation (CP1)
Environmental Information
Bioregions were mapped between 1999 and 2002 using expert knowledge and best available data and methods
30 reef bioregions 40 non-reef bioregions
Many bioregions previously lacked adequate protection
At least 20% of each bioregion included in a no-take zone
The GBR Marine Park

Non reef bioregions

Environmental Information
Other key issues:

Special and unique places
Critical habitats such as turtle nesting sites
Deep & shallow water sea-grass, fish spawning sites etc.

Special and unique places
Critical turtle nesting areas
Environmental Information
Biophysical Principles guided selection and use of environmental information
The Principles :
were developed by independent reef scientists
published in CP1
said that at least 20% of each bioregion had to be in no-take zones
No-take zones must be
arranged to form viable network, allowing connectivity, provides insurance policy

Social & Economic Information
Recreational fishing diaries, and tag and release records
Commercial fishing log-books
The location of boat-ramps and coastal developments
Historic ship-wrecks
Visitor use data

Over 10,000 submissions received in Phase 1 & >21,000 in Phase 2
All submissions read to identify community issues
All submissions were taken into account
Recreational fishing sites
Commercial fishing values
Using Social Information
Social, Economic, Cultural and Management Principles were:
developed by an independent panel of experts
published in Community Phase 1

The SEC Principles attempted to
minimise impact on existing users of the Marine Park
be fair—ie not impacting on one group or community more than another
but needed a Zoning plan easy to enforce
Previous Zoning
Previous Zoning, plus Trawl Plans
New green zones—environmental data only
Green zones—using economic data too
Green zones—revising boundaries
The Plan
What Does This Plan Do?
Provides strong, medium and long-term protection for future generations
Green zones mean more and bigger fish
Green zone spill-over, better fishing for reef communities
Natural values which attracts tourists and $ will be maintained
Protects at least 20% of each bioregion, special and unique areas, important habitats, and nesting areas—over 33% achieved

Phases of RAP
Classification (map biodiversity)
Reviewed existing protection
informal consultation with user groups
formal Community Participation phase 1
Identification of possible network options
Selection of most acceptable network
Draft zoning plan
formal Community Participation phase 2 (over 21,000 submissions)
Ministerial & parliamentary approval March 2004
Implemented July 1st 2004

Representative Areas Program
A new and effective network of ‘no-take’ areas representative of all bioregions helps to:
maintain biological diversity
maintain ecological processes and systems
provide an ecological safety margin, and if necessary, enable species and habitats to recover
ensure viable and sustainable industries
Current Status
Distribution of information and many maps to fishers, tourist operators, dive, boat and bait shops
Revised maps at boat ramps
Sorting out current permits in relation to new zoning, research stations issuing permits
Working with GPS manufacturers to incorporate zoning plans into charts, some available
Website available to download zoning plans for particular areas of interest

Related Activities
Reef Water Quality Protection Plan-implemented
Fisheries related: Reduction of number of fishing boats
Reduction in areas where trawling allowed
compensation being paid
Increased surveillance, penalties imposed
Dugong protected areas and reduce netting areas
Qld zoned adjacent coastal parks
Recognition of RAP
Authority awarded a Eureka Prize for Biodiversity Research and Banksia Environmental Award
WWF Australia acknowledges its importance for conserving biodiversity
Recognition overseas of importance of this approach to marine park management
Best practise
Relevance to Other Areas
Zoning with scientific basis
Problems facing the GBR faced by all reefal areas
Methods for zoning multi-use parks relevant to all areas in Australia and elsewhere
Such community involvement results in ownership and stewardship of the reef– schools adopting reefs, communities becoming effective policers
Other Management Strategies
Reef Water Quality Protection Plan
being implemented but ongoing and results will take years to be apparent
Reduction in number of fishing licences, compensation being paid
Increasing policing and enforcement
Global warming— the big question
increased rates of bleaching
increased cyclones activity
What is the long term future for the GBR?

Points to consider for Okinawa/Ryukyu Islands:
Only three major governments (National, 2 Prefectural).
However, management is very ambiguous.
Local fisheries have strong power; no no-take zones anywhere in Japan, aquaculture common.
Competing interests within national government have different agendas (Construction, Environment).
National laws for parks weak.
Okinawa Prefecture likely has strong wishes, but needs money from National government.

December 28th, 2011 class

Video presentation of coral reef biodiversity (in English).

December 14th, 2011 class

• 1. Review of evolution.
• 2. Introduction to reticulate evolution.
• 3. Examples from plants and fish.
• 4. Examples from corals.
• 5. Examples from zoanthids.
• 6. Conclusions
Part 1 - Evolution

Genetic Diversity
• Required to adapt to change in environment.
• Many methods of measurement.
• Large populations of naturally breeding animals have high genetic diversity.
• Reduced populations are concern.
Cnidaria DNA
mitochondrial DNA (mt DNA)
• evolves very slow in Cnidaria, opposite to most animals.
• 他の動物と違い、刺胞動物で進化が遅い。
DNA amd phylogenetics: All cells contain DNA - the code or blueprint of life.
This code has only four different “letters”: A, G, C, T.
Usual length 105 to 1010 base pairs.
生き物のひとつの細胞にある遺伝子の長さは105 to 1010 。
Genome projects read everything in one organism, but takes time and expensive.
Many studies use one or a few “markers” to investigate relations.

• By collecting the same marker from different samples and then analyzing them, we can make a tree.
• いくつかのサンプルから同じマーカーを読んで、並べてから、解析し系統樹を作る。
• It is thought/hoped a tree is similar to how evolution occurred.
• 系統樹から進化が見えると思われる。
Part 2 -
Reticulate Evolution
What is evolution?
• The descent of all organisms from a common ancestor.
• 全生物は共通の祖先から。
• The development of unique traits in response to environment, etc.
• 環境の変化などのせいで、それぞれのグループがユニークな特徴を持つ。
• Groups gradually “drift” away from each other.
• それぞれのグループが他のグループからだんだん離れる。
• But…
Some problems…
• How can “mega”-diversity arise?
• 非常に高い多様性はどうやって進化した?
• Even allowing for rapid evolution, there are cases of “mega”-diversity in very new and small environments, with many species adapted to very specific niches (plants, cichlids etc.).
• 時として、新しい環境で、種の数が想像以上に多い。
• Often hard to accurately explain “species” over large geographic scales.
• large geographic scaleで、種の説明や分類が困難になる場合がある。
• How can hybridization between species be explained?
• 別種のhybridizationも説明がしにくい。
Theory of evolution over time
• Evolution is evolving.
• Darwin - classic model.
• Currently, reticulate evolution is a “rare nuisance”.
• Likely our ideas will develop into an even more complex model.
Reticulate evolution?
• The pattern of evolution resulting from recombinational speciation.
• 種類Aと種類Bのハイブリッドによる進化。
• Not generally expected to be a common occurrence, but can explain “mega-diversity” in new environments and unexpected genetic results.
• 普通の進化より珍しいが、新しい環境などでは起こる可能性がある。
• Results in retainment of ancestral patterns in the genome, with “repackaging”.
• 遺伝子の配列は進化(変異)しない。ただ新しい組み合わせができるだけ。
• Believed to occur in many plant groups, and cichlids (fish).
• 植物やアフリカの池の魚類で起こっていると思われている。
Evidence of reticulate evolution
• Without laboratory experiments very hard to infer, but some ways:
• 研究室の実験以外で網状進化をどうやって見つける?
• Shared sequence portions between or within species.
• 種内、また種間の配列を見て、同じ部分があるかどうか?
• Differences between mitochondrial and nuclear DNA.
• ミトコンドリアDNAと核DNAの解析結果が違うかどうか?
Part 3 - Examples of Reticulate Evolution: Plants and Fishes
Example 1: peony flowers
(Sang et al. 1995)
• Sequenced ITS-rDNA of 33 species of Paeonia from Europe and Asia.
• Shrubs and herbs in northern hemisphere.
• Spotty distribution.
• Examined ITS-1 sequences.
• Many species showed additive patterns.
• Subsequent evolution has taken place in some species.

• Many hybrid species Asian.
• Parents of these hybrid species European.
• Suggests hybridization occurred in past.
• Can see historical patterns, useful in species with no fossil history.
• This type of evolution may be common in plants.
• In such cases must be careful with phylogenetics.
Another example:
Cameroonian crater
lake cichlid fish
• Megadiverse group of fish with monophyletic origin.
• Much research shows reticulate evolution may occur when nuclear and mt DNA phylogenies do not match.
• Invasion of new environments could trigger hybridization between species.
• Do hybrid swarms result from large areas with different environments or not?
• Cichlid fish provide great test case!
Barombi Mbo Lake
• 2.5 km in diameter.
• 110 m deep, only oxygen to 40 m.
• Four endemic genera; seven species.
• All on IUCN Red List - critically endangered.
• Evolved over 10000 years.
Materials and methods
• Two mt DNA markers and 2 nuclear markers.
• All types of fish from lake sampled; specimens deposited in museums.
• Differences in mt DNA and nuclear DNA.
• Secondary hybridization after evolution.
• Two ancient lineages formed new species; Pungu madareni.
• Hybrid speciation can make complex species assemblages even without prior hybridization.
Part 4 - Examples of Reticulate Evolution: Corals
Reticulate Evolution in Cnidaria?
• Several studies hint at reticulate evolution in Cnidaria, particularly corals and related groups.
• 特に花虫綱で網状進化の可能性がある。
• Marine environments where coral reefs are found are generally “new”.
• サンゴ礁の環境は比較的新しい。
• Centers of “mega-diversity” with “hyper-evolution” to micro-niches.
• 狭い地域で、多様性が非常に高い。
Acropora spp.
(Odorico & Miller 1997)
• Acropora very diverse, much morphological variation.
• Hybridization known from lab tests.
• ITS-rDNA shown to be a useful tool to detect this.

• Six colonies from five species.
• 18S rDNA and 28S rDNA obtained as well as ITS-rDNA.
Acropora ITS rDNA very short.
• Unexpected patterns of diversity, even within individuals!
• Such patterns consistent with ongoing reticulate evolution.
• Much more diversity than seen in plant ITS-rDNA.
• Could be due to more hybridization over longer ranges.
• Hybridization may occur over biological (not geological) time scales.
More corals
(Vollmer & Palumbi 2002)
• Examined all three Caribbean Acropora spp.
• Examined 2 nuclear and one mt DNA marker.
• A. cervicornis and A. palmata distinct species.
• A. prolifera are F1 hybrids.
• Shape of A. prolifera depends on which species provided egg.
• F1 hybrids are immortal mules that may occasionally hybridize.
• Hybrids may be common in corals.
Part 5 -
Reticulate evolution in zoanthids

Zoanthus spp. according to mt COI DNA
mt COIの結果による、マメスナギンチャク属の多様性
• Three species found with varying distribution. All ecologically similar to hard corals.
• 3つの種。生態はイシサンゴと似ている。
• Clear morphological variation between all three species.
• それぞれの種を区別できるようになった。
• This appears to be normal evolution.
• このデータから、普通の進化が推測できる。
• All Z. kuroshio and Z. gigantus sequenced as expected.
Z. kuroshioZ. gigantusの結果はそれぞれが単系統。
Z. sansibaricus had unusual results.
• 一方、 Z. sansibaricusの結果は単系統ではなかった!
• Some (2/3) samples gave expected sequences.
• 2/3のサンプルの配列(sansi)はmt DNAでの系統的位置と同様だったが、
• Some samples had both expected sequences and unknown “B” sequences.
• いくつかのZ. sansibaricus は不思議な “B”配列と普通の配列(sansi) 、両方を持つ。
• Some samples had only “B” sequences.
• 残りのZ. sansibaricus は不思議な “B”配列しか持っていない。
• B is closely related but different than Z. gigantus.
• “B”はZ. gigantus と近縁である。

Zoanthus undergoing reticulate evolution?
• Samples with normal sequences and with normal/B, or just B have normal Z. sansibaricus morphology.
• 全てのZ. sansibaricusの形態が同じだった。
• Could B-only be F2 - resulting from backcrossing or F1 x F1 crossing?
• “B”配列しか持っていないサンプルはF2?
Z. sansibaricus mass spawns, same as coral. No distribution barriers.
• マメスナギンチャク類はサンゴの様に同時に産卵する可能性がある。
• COI and morphology suggests NOT incomplete lineage sorting.
• 形態の結果やmt DNA配列を見ると、 incomplete lineage sortingじゃないと思うことができる。

Possible scenario for Zoanthus evolution
• Ancestor of Z.gigantus/B underwent one way hybridization (male B X female sansi), introducing B allele into Z. sansibaricus species.
Z.gigantus/Bの精子(nuclear DNA)がZ. sansibaricus 種内に入ってきた。
• Modern-day Z. sansibaricus has both B and sansi alleles, ancestral B/giga evolved into modern Z. gigantus.
• 現在のZ. sansibaricusはsansiもBも持っている。
• 現在のZ. gigantusは昔のZ.gigantus/Bから進化した。

More zoanthids
(Reimer et al. 2007b)
• Investigated Palythoa spp. in Japan.
• Thought to be two genera, but mt DNA shows one genus.
• P. tuberculosa and P. mutuki very closely related.

• ITS-rDNA shows two species (P. tuberculosa & P. mutuki) very closely related.
• Some specimens with intermediate morphology also apparently intermediate in phylogeny.
Results (2)
• Alignment of ITS-rDNA shows “reticulate” patterns between intermediates of two species.
• Appears as if some P. tuberculosa DNA has entered into P. mutuki population.

Conclusion 2
• In the future, more reticulate evolution will be found.
• This will impact conservation and our understanding of species.

Conclusion 3
• This will lead to better understanding of other related evolutionary events, such as lateral gene transfer (LGT).
References cited:
1. Sang et al. 1995. Documentation of reticulate evolution in peonies (Paeonia) using internal transcribed spacer sequences of nuclear ribosomal DNA: Implications for biogeography and concerted evolution. PNAS USA 92: 6813-6817.
2. Schliewen & Klee. 2005. Reticulate sympatric speciation in Cameroonian crater lake cichlids. Frontiers Zool 1:5.
3. Odorico & Miller. 1997. Variation in the ribosomal internal transcribed spacers and 5.8S rDNA among five species of Acropora (Cnidaria; Scleractinia): Patterns of variation consistent with reticulate evolution. Mol Biol Evol 14: 465-473.
4. Vollmer & Palumbi. 2002. Hybridization and the evolution of reef coral diversity. Science 296: 2023-2025.
5. Reimer et al. 2007a. Molecular evidence suggesting interspecific hybridization in Zoanthus spp. (Anthozoa: Hexacorallia). Zool Sci 24: 346-359.
6. Reimer et al. 2007b. Diversity and evolution in the zoanthid genus Palythoa (Cnidaria: Hexacorallia) based on nuclear ITS-rDNA. Coral Reefs 26: 399-410.
7. Shiroma and Reimer 2010. Zoological Studies.

December 7th, 2011 class

1. Review of dangers facing coral reefs (bleaching!).
2. How to stop bleaching (?).
3. Red coral in the Mediterranean.
4. The importance of fish and mangroves to coral reefs.
5. Community conservation in the Philippines.
6. Conclusions.
Part 1 - Review of dangers facing coral reefs (bleaching!)
Dangers facing coral reefs
Global warming is raising the temperature of the ocean; this kills corals - “coral bleaching”.
Also, as the oceans become more acidic, it is more difficult for corals to make their skeletons.
Perhaps 90% of coral reefs will be dead by 2050.

Crown-of-thorns starfish outbreaks

Dynamite and cyanide fishing

Coral bleaching: Images from Phuket, Thailand 2010
Corals (and many other coral reef invertebrates) are in symbiosis with Symbiodinium (zooxanthellae).
This symbiosis allows these invertebrates to live in nutrient-deficient sub-tropical and tropical waters.
Algal-animal symbioses are a successful strategy that has been repeated many times in evolution.
Weak point
Despite the success of this symbiosis, it has one very serious weak point:

Symbiodinium are very sensitive to low and high temperatures.

<18°C, and >30°C.
Coral bleaching
When temperatures are abnormal for the holobiont, stress occurs.
With this stress, thylakoids in Symbiodinium begin to break down; the symbiont begins to poison the host.
Corals lose their symbionts, either through cell-death, or by expelling them.
Hosts turn white = coral bleaching.

Predicting coral bleaching
The NOAA (USA) has spent much time on predicting bleaching.
Can now predict bleaching very accurately.
These tools available for free on the internet.
SST=sea surface temperature
DHW=degree heating weeks
Daily max=expected average maximum SST for a certain day
MMM=maximum monthly mean, average temperature of the hottest month
SST anomolies
Observed SST – daily max SST
Can be used to see what location is hotter than usual.
Coral bleaching HotSpot
Predicts what areas have thermal stress that can cause/contribute to coral bleaching.
HotSpot=observed SST - MMM
Degree heating week (DHW)
However, it is not just anomolies and hotspots that cause coral bleaching.
The total stress from the past weeks is important. One hot day does not kill most coral!
DHW=0.5*(sum of previous 24 HotSpot reports), where HotSpots <1.0°C are not counted.
Example: 1 week of 2°C higher than normal = 2 weeks of 1°C higher than normal.
DHW >8.0 usually can cause coral death.
Thailand’s situation
Very high DHW on both sides of the Malaysian Peninsula.
Made worse by no cold temperatures last winter.
Also, lack of wind (“doldrums”) causes more solar/UV stress, which makes bleaching worse.
Ko Racha

Ko Tao
Ko Samet
Thank you.
Part 2 -
How to stop bleaching (?)
West & Salm 2003
What factors help corals against bleaching?
Reviewed all research up until 2003.

Many examples of resistance to or recovery from bleaching.

Many factors contribute to resistance.
Can be included in management plans.
Cumulative stresses worse than one stressor.

Part 3 - Red coral in the Mediterranean
Red coral
Corallium rubrum is a precious coral in the Mediterranean.
Found 10 -250 m.
Harvested for long time, over-exploited.
Harvest reduced 66% in last 15 years.

Population structure
Two population types, large deep colonies and shallow small colonies.
Large drop off in shallow water at age 4, due to sponges and collection.

Genetic distance becomes significant at 100s of kms.
Thus, preservation of numerous populations needed.
Management on regional scale needed.
Must avoid local extinctions.
Conservation recommendations
Must be managed at national and international scales.
Only policy that works for such species.
Set minimum colony sizes, maximum yield per area, harvesting seasons.

Mumby et al. 2004
Reef fish often use mangroves as nurseries.
But can use other environments, not confined.
Also, despite deforestation, other pressures (fishing, larval supply) likely larger.

Management should include connected habitats, not islands of each type.
Future destruction of mangroves will have negative influence on reef.
Mumby et al. 2007
Caribbean reefs have damage from loss of Diadema antillarum and two species of coral.
“Sick” reefs characterized by macroalgae.
Can macroalgae be reversed? Or is it a stable state?

Used computer modeling and simulation.
Showed reefs can easily change to other states once D. antillarum died off.
With only parrotfish as grazers, small negative change in parrotfish numbers results in macroalgae blooms.
Coral becomes unstable state with low grazing.

Regular impact of hurricanes worsens with lack of grazers (fish and urchins).
Modeling useful for conservation targets.
Part 5 - Community conservation of coral reefs
Philippines consist of 7000+ islands.
Centuries have used reefs for livelihood.
Since 1970s, threatened by over-exploitation and destructive fishing methods.

Conservation started in 1974. Many projects failed.
Politics tied to conservation.
Local governments have authority but not knowledge or budget.
To be successful, combination of local and national people.
Within local group, must include users of reef; fishermen, resort owners, coastal residents, scuba divers.
Start of conservation
MDCP started in 1986 on three islands (62-166 households); Apo, Pamilacan, Balicasag.
All had less fish catch, increasing destruction and poverty.

MCDP plan
Marine reserves with buffer areas to increase number and diversity fish.
Development of local knowledge and alternative work.
Community center.
Outreach and replication program.
MCDP steps
Integration into community.
Education - marine ecology and resource management.
Group building, formalizing, strengthening.
Apo & Pamilacan remain strong.
Balicasag protection groups somewhat weakened due to large PTA resort and less local “ownership”.
PTA has good points too.
All islands have stronger municipal laws now.
Local fisherman believe sanctuary has helped.
Comparison of 1985-86 data with 1992 shows increases in fish, stable coral cover.
MPAs work on small islands by preventing destructive fishing and making locals understand value of conservation.
Small islands easier to implement plans.
Immediate benefits must be seen.
Baseline data necessary.
Local fishermen help with MPA location decisions.
Locals must understand how problem and answer related.
Management groups must have respected members.
Link with all potentially helpful groups.
All plans vulnerable to politics and outside groups.
Part 6 - Conclusions

Towards the future
In the future, more conservation plans will be implemented.
The gap between well protected areas and those not protected will widen.

Towards the future
Very few non-protected reefs will survive.

References cited:
1. West & Salm. 2003. Resistance and resilience to coral bleaching. Conservation Biol 17: 956-967.
2. Santangelo & Abbiati. 2001. Red coral: conservation and management of an over-exploited Mediterranean species. Aquatic Conserv Mar Freshwater Ecosys 11: 253-259.
3. Hughes et al. 2002. Biodiversity hotspots, centres of endemicity, and the conservation of coral reefs. Ecol Let 5: 775-784.
4. Roberts et al. 2002. Marine biodiversity hotspots and conservation priorities for tropical reefs. Science 295: 1280-1284. 8. 5. 5. Mumby et al. 2004. Mangroves enhance the biomass of coral reef fish communities in the Caribbean. Nature 427: 533-536. 9.
6. Mumby et al. 2007. Thresholds and the resilience of Caribbean coral reefs. Nature 450: 98-101.
7. White & Vogt. 2000. Philippine coral reefs under threat: lessons learned after 25 years of community-based reef conservation. Mar Poll Bull 40: 537-550.

November 30th, 2011 class

1. Quick introduction to diseases.
2. Common coral reef diseases.
3. Why are diseases becoming common?
4. How do diseases affect conservation?
5. Terpios: a new threat
6. Conclusions
Part 1: Disease
Example 1: Plague in humans

• Plagues have struck humans many times.
• Often kill 10-50% of population.
• Caused by an influenza virus.
• Two most infamous cases are 13th century Black Plague, and 1919-1920 Spanish Influenza.
• No one knows where plagues came from.
• Spread through common routes of trade.
• Spread faster in modern cases.
• Often affects young adults worse due to “cytokine storms”.
Spanish Influenza
• In some countries fatalities were as high as 50%.
• Killed more people than WWI.

How does this happen?
• New mutation in influenza virus that most humans do not have capability to respond to.
• Genetic variation provides resistance.
• SARS is a more recent case.

Example 2:
Introduction of a new disease into an isolated area
Elm trees common in North America and Eurasia.
Preyed upon by two species of bark beetles.
Beginning in the 1910s, some elms began to die.
Die-offs became rapid in 1960s.
Bark beetles somehow involved in the disease.

Survival of elms close to 0%.
• The causative agents of DED are ascomycete microfungi.
• Carried by the elm bark beetles.

• Three species are now recognized: Ophiostoma ulmi, which afflicted Europe in 1910, reaching North America on imported timber in 1928, Ophiostoma himal-ulmi, a species endemic to the western Himalaya, and the extremely virulent species, Ophiostoma novo-ulmi, which was first described in Europe and North America in the 1940s and has devastated elms in both areas since the late 1960s.
• The origin of O. novo-ulmi remains unknown but may have arisen as a hybrid between O. ulmi and O. himal-ulmi.
Part 2: Common coral reef diseases
Introduction to
coral reef diseases
• Bacteria observed in corals in early 1900s.
• Diseases noticed in 1970s, seemingly increasing over last 30 years.
• 34 mass events, affecting sponges, seagrasses, cetaceans, urchins, fish, molluscs, corals.
• Have changed composition of reefs.

Diseases affecting Scleractinia
• Many diseases named, but very little known.
• Most pathogens still unknown.
• Most common in Atlantic (Green & Bruckner).
• Not to be confused with coral bleaching.
Green & Bruckner 2000
Black Band Disease (BBD) Caused by numerous cyanobacteria (500 spp.) as a microbial mat.
Mat makes the colored band.
First observed in 1973.
Moves 3mm to 1cm/day.
Found in 42 spp. of coral.

Kuta & Richardson 2002
• BBD correlates strongly with depth, temperature, nitrites.
• Also correlates with diversity and orthophosphate.
White band disease: Pathogen unknown, may be bacteria. Noticed in 1981.
Tissue loss from base to tip.
Affects two species, Acropora cervicornis and A. palmata.
Moves 3mm to 1cm/day.
• WBD has drastically altered Caribbean reefs.
• Shifts in coral species.
• Loss of overall coral cover; algae increasing.

• Both species now “threatened”.
• Losses of over 98% of A. cervicornis. Locally extinct.
White plague: Affects many species, but no acroporoids.
Caused by Aurantimonas bacteria.
First observed in 1977.

Aspergillosis: Caused by terrestrial fungi.
Affect mainly Atlantic gorgonians.
Also affects waterfowl.
Noted in 1997.

Tumors: Similar to cancer.
Affects mainly A. palmata.
Irregular growth, no zooxanthellae.
Noted in 1960s and 1970s.

Other diseases: Many other diseases.
Mostly known from Atlantic.
Yellow band disease, yellow spot disease, white pox disease, brown band disease.
Most noted for first time in last 20 years.
Pathogens usually unknown.

Part 3: Why are diseases becoming common?
1. Global warming?
• Many people blame global warming.
• But likely much more complex.
2. Nutrient enrichment - Bruno et al. 2003
• Experiments done with YBD and Aspergillosis.
• Controls were disease only, experimental with added nitrogen and phosphorus.
Results - Aspergillosis
• Nutrients increased severity of disease in sea fans.
Results - YBD
• Presence of nutrients increased rate at which YBD developed in two species of coral.
3. Dust? -
Garrison et al. 2003
• Airborne dust from Africa and Asia carries many contaminants to reefs.
• Global warming and desertification increasing dust, therefore increasing contaminants.

Part 4: How do diseases affect conservation?
Effects are widespread
Many studies have documented widespread coral decline in almost ALL coral species.
Porter et al. 2001 showed many declines 1996-1998 NOT due to coral bleaching but disease.

• Porter et al. 2001 cont
• Green & Bruckner 2000

• Green & Bruckner 2000
Many examples of diseases spreading, many examples of reef degradation (show many photos).

Overview of disease
• All diseases have negative effects.
• Only WBD has changed communities drastically.
• Pacific 15 years behind Atlantic.
• Compounded negative influences more severe for coral reefs.

Part 5: Preliminary results of field surveys of Terpios outbreaks in the Nansei Islands, Japan

Terpios hoshinota Rützler and Muzik 1993
Terpios in the Nansei Islands - history
Outbreak noticed in Mariana Is. 1973. (Bryan 1973 )
Terpios-Nansei project
Assess the current distribution of Terpios in the Nansei Islands.
Establish monitoring sites.
If present, characterize sexual reproduction & ecology.
Survey all major islands by snorkel/scuba (Reimer).
Monthly/bi-monthly sampling at designated locations – histology (Hirose), genetics (Chen).
Permanent transects at massive outbreak (Reimer), analyses (Reimer, Nozawa).
Preliminary results
Three situations observed: none, small amounts, massive outbreak
Yonama, Tokunoshima had massive outbreak (87.9% cover) in 1986 (Marine Park Center Foundation 1986).
Terpios absent or present in small numbers in most reefs in Nansei Islands (38/39 examined locations).
Coverage does not appear to fluctuate much in most locations.
Discussion & Questions
Massive outbreaks still occur in Nansei Islands.
How long do outbreaks last?
“Recovery” observed at Yonama, but is this true recovery? At least, not a dead-end.
Results suggest outbreaks are linked to reef degradation, but factors not clear.
Future work
Permanent transect results & analyses.
Try to quantify speed at which massive outbreaks can occur.
Combine analyses with genetic, histological results.
Examine Yakomo (current outbreak location) to understand causes of outbreaks. Why this location?

Part 6: Conclusions.
Conclusion 1
• Disease more widespread on reefs in Caribbean.
• More research? Partially.
• Monitoring in Pacific very critical.
Conclusion 2
• Only one disease has permanently changed community structure (WBD).
• Other diseases locally important.
Conclusion 3
• Very few studies have investigated in detail mortality rates.
• Monitoring of individual colonies needed.
Conclusion 4
• Diseases increasing.
• Bleaching appears to be more critical, but two problems appear related.
Conclusion 5
• Diseases not well understood.
• Many diseases affect many species; possibly more or less diseases.
• Pathogens need to be investigated.
Conclusion 6
• While bleaching currently more serious, foolish to ignore diseases.
• May be “indicator” of serious problems, similar to amphibians.
What needs to be done
• <3% of reefs in danger have low human impact.
• More research needed on human influences and pathogens.
• Management and conservation then follow.
1. Green & Bruckner. 2000. The significance of coral disease epizootiology for coral reef conservation. Biological Conservation 96: 347-361.
2. Aronson & Precht. 2001. White-band disease and the changing face of Caribbean coral reefs. Hydrobiologia 460: 25-38.
3. Garrison et al. 2003. African and Asian dust: from desert soils to coral reefs. BioScience 53: 469-481.
4. Bruno et al. 2003. Nutrient enrichment can increase the severity of coral diseases. Ecology Letters 6: 1056-1061.
5. Kuta & Richardson. 2002. Ecological aspects of black band disease of corals: relationships between disease incidence and environmental factors. Coral Reefs 21: 393-398.
6. Porter et al. 2001. Patterns of spread of disease in the Florida Keys. Hydrobiologia 460: 1-24.
7. Reimer, Hirose, et al. new Terpios papers.