[Coral-List] Coral species list for Mesoamerican Barrier Reef System

Szmant, Alina szmanta at uncw.edu
Tue May 3 22:05:19 EDT 2011

Hi Derek:

A few years ago Bob Ginsburg elected me as an "honorary geologist" because of my interest in the processes of reef building, so here goes my 2 cents worth with regard your definition of a coral reef only consisting of the coral dominated forereef.

A coral reef is much more than just the forereef framework.  The sediments that infill the cracks and crevices and troughs of the forereef are derived from bio-and physical erosion of coral skeletons and all the skeletal material from calcareous algae and calcifying non-coral inverts.  All of that accumulation and infilling is part of the reef building process.  The reef crest and reef flat and backreef lagoon are all considered zones of a coral reef.  The forereef dominated by hermatypic corals is only one of the many zones of the reef and not the only one that accretes over time.  Without the buildng of the reef crest-back reef-lagoon, the forereef would have no support and not be able to keep up with sea level.  If you examine the nice review chapters by Hallock and Hubbard in Life and Death of Coral Reefs (1997; C.E. Birkelan editor) you will see clear explanations of the reef building processes and the variety of processes, organisms in addition to corals, that are critical to reef building.  In fact, in Hubbard's chapter, take a look at the figure of reef structure and fate with changes in rate of sea level rise.  The process of backstepping, and eventually drowning are the result of carbonate secretion not being able to keep on building the leeward parts of the coral reef if sea levels rises too fast.

As coralo-centric as I am, I recognize the crtitical contribution of all the calcareous algae of all colors to the reef building process.  Except for maybe some A cervicornis haystacks, and some the E Pac Pocillopora reefs, neither of which have much real framework, you won't find a coral reef (defined as the whole structure as commonly diagramed from head to toe, not just the forereef) that has more than ca. 50 % of the CaCO3 made up of coral skeleton.



Dr. Alina M. Szmant
Professor of Marine Biology
Coral Reef Research Program, Center for Marine Science
University of North Carolina Wilmington
5600 Marvin K. Moss Lane
Wilmington NC 28409
Tel:  US +1  9109622362  Call ; fax: (910)962-2410;  cell:  US +1  9102003913  Call
From: Derek Manzello [dmanzello at rsmas.miami.edu]
Sent: Tuesday, May 03, 2011 9:29 PM
To: Douglas Fenner
Cc: Szmant, Alina; coral-list at coral.aoml.noaa.gov; Peter Vroom
Subject: Re: [Coral-List] Coral species list for Mesoamerican Barrier Reef System


What you said is this: "That said, most reefs are not just
coral reefs, they are coralgal reefs or even algal coral reefs, with
coralline algae and other calcareous algae contributing as much or more
calcium buildup than the corals."

When you say 'most reefs', I interpret this to mean the actual reef, which
is the 3-D wave resistant structure.  There are sediment aprons that flank
all reef framework structures.  These are where the numbers come from that
Alina cited.  Often, a lot of this sediment is derived from other
calcareous organisms.  /Halimeda/ can contribute significantly to these
sediments.  This is essentially sand.   This is not the reef.  The reef
structure is built by corals, except for a few rare cases where you have
algal ridges that are frameworks constructed by crustose coralline algae
(CCA).  However, this is not the rule, but the exception, as many of these
CCA algal ridges actually are just coating the very outer veneer of
antecedent coral framework below (i.e., "standing on the shoulders of
coral reef frameworks").

When you look into the subsurface of the reef structure it is almost
always predominantly coral.  This is what Macintyre is pointing out.  His
point is that, for over a century, people have been swimming around and
making qualitative observations like "Look! That reef has a lot of CCA on
its surface, therefore CCA must have built it".  When you look at the
actual subsurface reef cores, rather than what sits or coats the surface
at any given point in time, its predominantly coral.  Just because a reef
may have high percent cover of CCA or something else, does not mean it was
built by CCA, except where wave energy is so high that corals can't live.
However, keep in mind that this high wave energy is usually only achieved
after corals have built the structure up to sea level, creating this niche
for CCA to dominate.

/Halimeda/ does have very high rates of carbonate production (grams CaCO3
per unit area per unit time).  However, this ends up as sediment, filling
in lagoons, depressions where it doesn't get washed away, or, in some
cases, cracks and crevices of the actual reef structure.  However, it is
incorrect to say that this is contributing to the build up of coral reefs
When you said buildup, I interpret this as construction of structure, not
the production of sand.  In a lot of cases, what makes this 'buildup' of
non-coral calcareous sediments possible is b/c corals have already built
the reefs that impede wave action/turbulence, allowing this to settle out
and accumulate on the reef perimeter.


----- Original Message -----
From: Douglas Fenner <douglasfenner at yahoo.com>
Date: Tuesday, May 3, 2011 7:55 pm
Subject: Re: [Coral-List] Coral species list for Mesoamerican Barrier Reef

> Derek,
>      My original statement was "coralline algae and other
> calcareous algae contributing as much or more calcium buildup than
> the corals."  People may well think that coralline algae are
> critical to coral reef structure, and your quotes say that's not
> true, and I would agree, there are surely reefs with very little
> crustose coralline algae.  But my statement didn't say that
> coralline algae are critical to reef structure, I carefully omitted
> that, and instead said that calcareous algae often contributed as
> much or more calcium buildup than corals.  Halimeda is a major
> producer of calcium, and my impression is that sand, partly from
> Halimeda flakes and partly from ground up coral, fills in spaces
> between hard coral skeletons, eventually being cemented together
> into solid reef.  "Calcareous algae" includes crustose coralline
> algae, other coralline algae, other red calcareous non-coralline
> algae (like Peysonellia), Halimeda, other green calcareous
> algae, and brown calcareous algae.  And there are other components
> contributing calcium like foraminifera and other invertebrates, as
> Alina points out.
>      When you refer to the sediments, I remind that the sediments
> are primarily calcareous, and they are primarily produced by
> organisms on the reef (though in some places there are surely major
> contributions from algae like Halimeda living on the sand itself).
> It is surely a reef-associated feature and in most cases reef-
> produced feature, and most people would consider it part of the
> reef system.  If you are actually on a reef, you will often notice
> that it is a mosaic of hard structures and soft, and it is often
> hard to say where one ends and another starts, in the sense that
> one area has lots of corals and hard substrate, and it fades into
> less and less corals and hard areas and more sediment until you
> reach a place which is mostly sediment.  Ridges and spurs are hard,
> the floors of grooves may be sand, holes filled with sand are all
> over, sand is often on top of reef rock, etc.  Sand produced by
> reefs is an integral part of reef systems and
> Halimeda a major sand producer.
>       As for the critical role corals play in producing a wave-
> resistant structure, I agree.  But I will point out that on algal
> ridges, the waves are so powerful that nothing but crustose
> coralline algae are wave-resistant enough to live there.  Corals
> are not there because they aren't sufficiently wave-resistant.  So
> while it only happens where the waves are the largest, and only in
> shallow water, and it is only a small part of a whole reef, this is
> a place dominated and built entirely by crustose coralline algae,
> which are the most wave-resistant of all reef organisms.
>       I will also quote the Steneck paper.  "Calcareous algae are
> important to reefs both today and in the past."  "Calcareous green
> algae are a major infiller of reefs.  Given their ubiquity and
> productivity they comprise a surprisingly high proportion of the
> total carbonate budget of reefs."  "Encrusting coralline algae can
> be a major space occupier on reefs."  "Coralline algae is not
> necessary for reef growth although under conditions of intense and
> frequent wave action at or near mean low water, significant
> coralline build ups, or algal ridges can develop."  "Crustose
> coralline red algae and calcareous green algae, such as Halimeda,
> dominate certain reef zones."  I would add that coralline algae is
> not totally restricted to shallow areas, we have a bay in American
> Samoa that gets quite a lot of wave surge, and not only is the
> benthos at 5 m deep dominated by crustose corallines, but there is
> nearly as much cover of it in places at 30 m
> deep.  That is unusual, but it can happen.  Also, there is a lot
> of crustose coralline algae and/or Peysonellia on slopes that is
> hidden under turf and macroalgae.  Crustose corallines and/or
> Peysonellia form an understory that is not as obvious as in shallow
> water, there is more there than it appears at first glance.
>      May I commend to readers a recent article available online
> open access:
> Vroom, P.S.  2011.  "Coral Dominance": A dangerous ecosystem
> misnomer?  Journal of Marine Biology, Volume 2011, Article ID
> 164127, 8 pages.
> This article supports this line of thinking with quite a bit of
> evidence, and providing quotes like  "...other healthy reef
> ecosystems were found to rely almost
> entirely on calcified algae and foraminifera for calcium carbonate
> accumulation."  and  "Calcareous algae include not only crustose
> coralline red genera, but also calcified macroalgae such as species
> of the segmented, green genus Halimeda, which are the main
> producers of carbonate sediments in many reef systems [16, 49–52]."
>   "In fact, in the least impacted Pacific ecosystems monitored by
> the National Oceanic and Atmospheric Administration, Pacific
> Islands Fisheries Science Center’s Coral Reef Ecosystem Division
> (CRED), the vast majority exhibit average island-wide percent cover
> of calcified red algae ranging from 1% to 42% and scleractinian
> corals ranging from 2% to 40% (Figure 2) [39]."
>       So while I would agree that crustose coralline algae are not
> critical for reefs, I would also say that calcareous algae as a
> whole are a very important part of reef ecosystems and the
> production of calcium, and can at times produce as much or more
> calcium than corals produce.     Cheers,  Doug
> ________________________________
> From: Derek Manzello <dmanzello at rsmas.miami.edu>
> To: szmanta at uncw.edu; coral-list at coral.aoml.noaa.gov
> Sent: Monday, May 2, 2011 2:48 PM
> Subject: Re: [Coral-List] Coral species list for Mesoamerican
> Barrier Reef System
> Hi Alina and coral-listers,
> Given the values you provided, I'm assuming you've based your ideas on
> Table 1 from Orme (1977) within v.4 of the Jones and Endean books.
> The numbers you refer to are from lagoon and reef sediments, not
> actualreef frameworks.  The title of the Table is, in fact, "Some
> examples of
> the composition of peripheral reef sediment".
> Nobody is arguing that there other calcifiers on coral reefs. A lot of
> these calcaerous materials end up in sedimentary environments,
> sometimesin domineering fashion (e.g., /Halimeda/ in some
> lagoons).  However, these
> sedimentary environments usually occur in the lee of A) structural
> coralreef framework or B) some other baffling antecedent topography.
> When I think of a coral reef, I think of a calcareous, 3-D, wave
> resistantstructure.  I do not consider the sediments that
> accumulate in the lee of
> structural reef frameworks the actual coral reef.  Based on your
> argument,any calcareous sedimentary environment should therefore be
> a coral reef.
> Saying these other calcifiers contribute to the building of coral
> reefs is
> like saying that the windows on a skyscraper are instrumental to its
> construction.
> I stand by Ian Macintyre's (1997) conclusion:
> "Crustose coralline algae do not contribute significantly to reef
> framework except in shallow water, high-energy environments.  In
> Holocenereefs, which have had to cope with advancing sea levels,
> these conditions
> have occurred mainly in the latter stages of reef growth, when
> there is
> little space left for framework accumulation.  Thus the coralline
> algalcontribution to the overall structural framework in modern
> reefs has for
> the most part been minor."
> Best,
> Derek
> Refs cited
> Macintyre IG (1997) Reevaluating the role of crustose coralline
> algae in
> the construction of coral reefs. Proc 8th Int Coral Reef Symp 1:725-
> 730
> Orme GR (1977) Aspects of sedimentation in the coral reef
> environment. In:
> Biology and Geology of Coral Reefs, v. 4.  Jones OA, Endean R (Eds),
> Academic Press, p. 129-176
> --------------------------------------------------------------------
> -------
> Hi Derek:
> Contrary to your statements, and those cited from the 8th
> InternationalSymposium, there are a number of studies that have
> quantified the relative
> contribution of the various calcifying taxa to reef building, and
> thereare large regional and local differences in these values.  In
> chapters in
> the old (1973-1975) Geology and Biology of Coral Reefs and other
> sources,analyzing which taxa make up how much of long reef cores,
> corals are
> attributed with 5-45 % of the carbonate, red and green calcified algae
> together 5 to 60 % of the carbonate, forams 2-40 %, and miscellaneous
> inverts (echinoderms, mollusks, crustaceans up to 30 % of the
> carbonate.
> So while I agree that the large coral heads and also the fast growing
> branching corals are essential for reef formation, they don't do it
> alone.
> Best,
> Alina
> *************************************************************************
> Dr. Alina M. Szmant
> Professor of Marine Biology
> Center for Marine Science and Dept of Biology and Marine Biology
> University of North Carolina Wilmington
> 5600 Marvin Moss Ln
> Wilmington NC 28409 USA
> tel:  910-962-2362  fax: 910-962-2410  cell: 910-200-3913
> http://people.uncw.edu/szmanta
> *******************************************************
> -----Original Message-----
> From: coral-list-bounces at coral.aoml.noaa.gov
> [mailto:coral-list-bounces at coral.aoml.noaa.gov] On Behalf Of Derek
> Manzello
> Sent: Monday, May 02, 2011 1:15 PM
> To: coral-list at coral.aoml.noaa.gov
> Subject: Re: [Coral-List] Coral species list for Mesoamerican
> Barrier Reef
> System
> Hello Doug and coral-list,
> Thanks for the helpful information.  However, I found one statement in
> your message inaccurate.
> The statement I refer to is this: "That said, most reefs are not just
> coral reefs, they are coralgal reefs or even algal coral reefs, with
> coralline algae and other calcareous algae contributing as much or
> morecalcium buildup than the corals."
> This is a popular quip among certain reef ecologists that is not
> supported by any hard data, other than some very rare and extreme
> cases.  I felt it was necessary to respond to this because this
> misconception seems to be popping up everywhere and I think we need to
> remind ourselves that this was already debated and pretty much
> resolveda while back.  In fact, there was an entire session during
> the 8th
> International Coral Reef Symposium entitled "Are calcareous algae
> important to reefs today or in the past?"
> In the session summary, Steneck and Testa state: ""Calcareous algae
> havelong been assumed to be important and perhaps even necessary
> for reef
> development. If so, one might expect to find the rates of reef growth
> (or accumulation) correlating with the abundance of calcareous algae.
> Macintyre (1997) reviewed the association of coralline algae and reef
> development by reviewing  non-algal ridge reefs and 5 algal ridge
> reefsthat have been drilled all over the world. Macintyre made the
> simple but
> profound point that most of the worlds fastest growing coral reefs
> wererelatively devoid of coralline algae during that period of rapid
> growth... Coral reefs do not require corallines or any calcareous
> algaefor their formation. Internal strength of reefs is primarily
> augmentedby submarine lithification and not by coralline algal
> cementation as has
> often been asserted."
> CCAs are important to reef dynamics for a variety of reasons, but they
> do not significantly contribute to the calcium carbonate budget of the
> vast majority of coral reefs, nor do they 'cement' reefs, as has
> been a
> popular misconception for years.  For further information on this
> topic,I refer the list to the important papers within this session
> (all freely
> available online at reefbase), that are all too often overlooked.  The
> specific papers referred to here are referenced below.
> With regards,
> Derek Manzello
> Macintyre IG (1997) Reevaluating the role of crustose coralline
> algae in
> the construction of coral reefs. Proc 8th Int Coral Reef Symp 1:725-
> 730
> Steneck RS, Testa V (1997) Are calcareous algae important to reefs
> todayor in the past? Symposium summary. Proc 8th Int Coral Reef
> Symp 1:685-688.
> On 4/21/2011 4:41 PM, Douglas Fenner wrote:
> > >      Almost all reef-building coral species in the Caribbean
> have ranges
> > > throughout the Caribbean, since the Caribbean is a relatively
> smallbody of
> > > water (compared to the Indo-Pacific, for instance).  Most all
> of  the
> species
> > > have already been found in places like Belize, Cozumel,  Akumal,
> Cancun area,
> > > and so on, and some of the others may be there but  just
> haven't been
> found
> > > there yet.  There are a very few which have not  been found in
> the NW
> or W
> > > Caribbean at all, and might (might) not be  there (to prove
> they are
> not there
> > > is like trying to prove the null  hypothesis).  One that is pretty
> sure not to
> > > be there is Millepora  squarrosa.  It is only known from the
> southeast> > Caribbean, and reports  elsewhere are likely all
> errors.  Millepora
> complanata
> > > can look a bit like it, but if you look in the Humann book
> you'll see M.
> > > squarrosa is actually quite distinctive and easy to
> recognize.  A second
> > > species is Leptoseris cailleti, a small deep-water  species
> that is
> rarely
> > > reported anywhere.  Millepora striata is rarely  reported, but I
> reported it
> > > from Belize, so it is in the MesoAmerican  reef system.  There
> are a
> few other
> > > rarely reported or less well known  species that may or may not be
> there, such
> > > as Madracis senaria, Madracis  asperula, Madracis carambi and
> Poritesbranneri.
> > > The situation is  quite different with the azooxanthellate corals.
> How many are
> > > present  in an area is poorly known, probably because they are
> small and
> > > cryptic,  but they may be patchy as well, since they typically
> live in
> very
> > > specific habitats like cavern roofs that are searched less
> often and
> less
> > > completely than open habitats.  Also, their identification is
> not a
> trivial
> > > matter for most  of us reef biologists, most require sending a
> sampleto the one
> > > or two  people in the whole world who are experts on their
> taxonomy(I'm not one
> > > of them, Dr. Stephen Cairns at the Smithsonian is one, and can
> put you
> in touch
> > > with the others).
> > >        For the zooxanthellate species, you can find range maps in
> Veron (2000),
> > > but it appears he fills in all the Caribbean for any species found
> somewhere in
> > > the Caribbean.  He's working on a much more detailed  database
> called"Coral
> > > Geographic."
> > >
> > >       To my way of thinking Belize has a true barrier reef, but
> therest of the
> > > MesoAmerican reef system is not a barrier reef as far as I
> know, but
> I'm no
> > > expert on it.  A barrier reef has to have a significant lagoon
> betweenit and
> > > land, and my impression is outside Belize, reefs are pretty much
> fringing.  I've
> > > also heard of the Florida Keys reefs referred to as a barrier
> reef.  I
> prefer
> > > the older name, "Florida Reef Tract" since as far as I know it
> consists of a
> > > series of relatively small reefs with wide gaps between them,
> and more
> > > continuous ridges of hard grounds that are not currently living
> coralreefs and
> > > don't get close to the surface.  Gene Shinn also tells me that the
> Florida Keys
> > > reefs have been called bank reefs.  That said, most reefs are
> not just
> coral
> > > reefs, they are coralgal reefs or even algal coral reefs, with
> coralline algae
> > > and other calcareous algae contributing as much or more calcium
> buildup than the
> > > corals.  Also, the Great Barrier Reef is not a single reef but
> a whole
> series of
> > > about two  thousand reefs, with gaps of various sizes (a maze
> that in
> effect is
> > > a barrier to  navigation unless you have GPS and a very good map
> system and are
> > > a good  navigator).  There is one section that is a nearly
> continuousbarrier,
> > > the section called the "Ribbon Reefs."  I'd also remind people
> of the
> barrier
> > > reef in New Caledonia, which is like Belize and the Ribbon
> Reefs in
> the GBR, a
> > > nearly continuous barrier with some small gaps.  New Caledonia
> is said
> to have
> > > the longest continuous barrier reef in the world, and likely
> that is
> not widely
> > > known.  Anyhow, "MesoAmerican reef system"  sounds fine with
> me, as
> does Belize
> > > Barrier Reef, but adding barrier to  MesoAmerican does not, nor
> doesit for
> > > Florida.  It seems like today  people think the word "barrier"
> addscharisma, so
> > > they want to call  their reef a barrier reef.  Fringing reef
> ought to
> also have
> > > some  charisma, think of the Ningaloo fringing reef in western
> Australia,> > longest fringing reef in the world.  Not nearly as
> well known as the
> GBR, but a
> > > huge and amazing reef.  Think of Indonesia, which has more  coral
> reefs than any
> > > other country in the world (slightly more than  Australia), I
> bet most
> of their
> > > reefs are fringing.  Also among the most  diverse in the world,
> a true
> world
> > > treasure.  Fringing is good.        Doug
> > >
> > > Cheers,  Doug
> > >
> > > Fenner, D. 2001.  Biogeography of three Caribbeancorals
> (Scleractinia);> > Tubastraea
> > >
> > >     coccineainvades the Gulf of Mexico.  Bulletin of Marine
> Science 69:
> > > 1175-1189.
> > >
> > > Fenner, D.  1999.  New Observations on the Stony Coral Species
> (Scleractinia,
> > >     Milliporidae, Stylaseridae) of Belize(Central America) and
> Cozumel(Mexico).
> > >     Bulletin of Marine Science 64: 143-154.
> > >
> > > Fenner, D. P. 1993. Some reefs and corals of Roatan (Honduras),
> CaymanBrac, and
> > >     Little Cayman.  Atoll Research Bulletin 388: 1-30.
> > > Weerdt, W. H.  de.  1990.  Discontinuous distribution of the
> tropicalwest
> > > Atlantic  hydrocoral Millepora squarrosa.  Beaufort. 41: 195-203.
> > >
> > > Douglas Fenner
> > > Coral Reef Monitoring Ecologist
> > > Dept Marine & Wildlife Resources
> > > American Samoa
> > >
> > >
> > > Mailing address:
> > > PO Box 3730
> > > Pago Pago, AS 96799
> > > USA
> > >
> > >
> > > work phone 684  633 4456
> > >
> > >
> > > Sharply increased mass loss from glaciers and ice caps in the
> CanadianArctic
> > > Archipelago
> > >
> > >
> > > Between  the periods 2004-2006 and 2007-2009, the rate of mass
> losssharply
> > > increased from 31 ± 8 Gt yr 1 to 92 ± 12 Gt yr 1 in direct
> response to
> warmer
> > > summer temperatures, to which rates of ice loss are highly
> sensitive(64 ± 14
> > > Gt yr 1 per 1 K increase).
> > >
> > > Gardner et al Nature
> > >
> http://www.nature.com/nature/journal/vaop/ncurrent/full/nature10089.html?WT.ec_id=NATURE-20110421
> > >
> > >
> > >
> > >
> > >
> > >
> > > ________________________________
> > > From: Brittany Huntington <brittanyhuntington at gmail.com>
> > > To: coral-list at coral.aoml..noaa.gov
> > > Sent: Thu, April 21, 2011 4:13:17 AM
> > > Subject: [Coral-List] Coral species list for Mesoamerican
> Barrier Reef
> System
> > >
> > > I am interested in determining the regional species pool for
> scleractinian
> > > corals within the Mesoamerican Barrier Reef System.  Published
> reportsweigh
> > > in around 60 species from what I have found but would
> appreciate any
> leads
> > > to a taxonomic list of coral species observed in the region.
> > >
> > > Thanks in advance,
> > > Brittany Huntington
> > >
> > > Brittany Huntington
> > > Doctoral Candidate
> > > Division of Marine Biology and Fisheries
> > > Rosenstiel School of Marine and Atmospheric Science
> > > University of Miami
> > > 4600 Rickenbacker Causeway
> > > Miami, FL 33149
> > > _______________________________________________
> > > Coral-List mailing list
> > > Coral-List at coral.aoml.noaa.gov
> > > http://coral.aoml.noaa.gov/mailman/listinfo/coral-list
> > > _______________________________________________
> > > Coral-List mailing list
> > > Coral-List at coral.aoml.noaa.gov
> > > http://coral.aoml.noaa.gov/mailman/listinfo/coral-list
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