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Arizona and Arizona State are bitter rivals, but they also have a lot in common at present.
Both were severely disappointing in 2011. Both have new head coaches as a result. Those new coaches, Rich Rodriguez at Arizona and Todd Graham at Arizona State, know each other well. They both are somewhat controversial for various reasons. Both run up-tempo offenses that looked pretty good in week one. And both won.
Further, both will be home against major-conference foes on Saturday. Both teams lost on the road to those foes last year.
Of course, the parallels are not exact. Arizona is a double-digit underdog to No. 18 Oklahoma State, while Arizona State is a slight favorite over Illinois.
The Wildcats are playing the Cowboys for a third time in 15 months. Both previous meetings were blowout defeats. But these aren't the same Cowboys who nipped Stanford in the Fiesta Bowl. They lost quarterback Brandon Weeden and wide receiver Justin Blackmon and welcome back just 12 starters. Their new signal-caller, freshman Wes Lunt, will be making his first road start after receiving no challenge at all last weekend against a dreadful Savannah State team.
"They could have scored 184 by halftime if they'd kept their starters in," Rodriguez said of the Cowboys' farcical 84-zip victory.
Illinois beat the Sun Devils 17-14 last year, but those programs' seasons were notably similar. Both Arizona State and Illinois started fast, earned national rankings, then cratered. And, therefore, fired their coaches.
Another tie that binds: New Illinois coach Tim Beckman coached Toledo, the team Arizona beat last weekend, for three years before arriving in Champaign, losing at home to the Wildcats in 2010.
Still, while everyone will know each other in points central and south in Arizona, there are plenty of unknowns. Last season's game films shouldn't be too revealing for anyone.
"We watched it a whole bunch looking at the personnel," Graham said. "I think it helps us evaluate their personnel but their defense is different, their offense is different. It's all new coordinators so from a schematic standpoint it didn't help us at all. From a personnel standpoint, studying their personnel, that is what we mostly used it for."
And even that is a bit confusing. Veteran Illini quarterback Nathan Scheelhaase is decidedly questionable after hurting his ankle during a 24-7 win over Western Michigan. The Sun Devils might be facing a dual-QB system with Reilly O'Toole and Miles Osei.
That would be a big advantage for the Sun Devils, though Graham said he's preparing his team as if Scheelhaase will play. On defense, the Illini are stout. It should give Graham's up-tempo offense and new starting QB Taylor Kelly a far bigger challenge than Northern Arizona did in week one.
Still, the Sun Devils appear far more likely than the Wildcats to improve to 2-0 and generate further positive momentum for Graham.
The Wildcats have a puncher's chance for an upset, but their defense is going to have to catch a few breaks. The unit played fairly well against Toledo, at least based on low expectations entering the game. But Oklahoma State, the defending Big 12 champion, is not Toledo. Even with some new players in key positions, the expectations for a lot of points will continue in Stillwater. The Cowboys have been consistently good on offense under coach Mike Gundy.
"It's an opportunity for us to play a ranked team," Rodriguez said. "I told our team there's not a lot of people who talk about Arizona football in certain circles, and if you want to become more relevant, the best way to do it is to beat a ranked team. It's a great challenge but also a great opportunity."
Arizona and Arizona State are constantly trying to one-up the other, and it will be popular to compare the records Rodriguez and Graham produce their first seasons. While both are ultimately trying to build programs that consistently compete in the top half of the conference, the competition and rivalry will be present each step of the way.
So fans of both programs are hoping, with considerable zeal, that they don't share anything in common at the stroke of midnight on Saturday.


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subset: commoncrawl


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Comfort Socks Newsletter - May 2013
New Socks for Homeless Shelters
Your support of Comfort Socks has allowed us to receive 44,635 pairs of socks so far. We are so grateful to all our supporters. First: We wish Comfort and Compassion are known by our actions and to all sock recipients. Second: We pray for Blessings for our supporters, sock recipients, and those without a home. Third: Please stay mindful of us and remember us monthly, if you can. Special Thanks to New Life Christian Fellowship for allowing us the use of their facilities and for their prayers and support for our mission.New Life Christian Fellowship
Comfort Socks is a 501(c)3 public charity
that collects and distributes new
socks for homeless
people and families. All contributions are tax-deductible as allowed by law.


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Marshals players come from throughout the Metroplex and North Texas area.
Kelly Field
2000 Kelly Blvd, Carrollton, TX 7500
To register for a tryout, fill out a "Play for the Marshals" form HERE. For info, call 972-841-3744 or email firstname.lastname@example.com.
The Marshals 16U - Bern team, coached by head coach Brian Bern, won the NTX Southwest Premier Finale championship, capturing wins along the way against, DBAT, Stix, Blackhawks and Cavemen teams.
Former Marshals SS Nolan Ritchie (Frisco Centennial, Cisco JC) has signed to play with The University of Alabama.
Congratulations to the 2016 Marshals players moving on to the next level
Marshals players understand that academic success is a critical component of being a student-athlete.
We are very proud of these players who have worn the Marshals colors, and wish them well as they move on to the next level.
|Reese Allen
|Richland College
|Trevor Baxter
|Richland College
|David Clairmont
|Olney Central College
|Christopher Cloud
|Northern Oklahoma
|Blake Divalero
|Oklahoma Wesleyan University
|Peyton Graham
|Murray State
|Jake Hillebrenner
|Centenary College
|Levi Hon
|Midland College
|Matt Horvath
|Spring Hill College
|Brady Howle
|UT Arlington
|Brett Hudspeth
|Hardin Simmons
|Jared Kengott
|Southern Illinois University
|Henry Mikkelson
|Aquinas College
|Damarco Perkins
|Tyler Junior College
|Patrick Roseborough
|Cloud County CC
|Coleman Sanford
|Wayland Baptist University
|Gage Scribner
|Murray State
|Ford Sherrington
|Hendrix College
|Seth Spears
|Texas Wesleyan University
|Zach St. Pierre
|Eastfield College
|Jon Urso
|East Texas Baptist
|Logan Worswick
|McPherson College


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HBO Buzz
Woe betide anyone trying to cut ahead of Larry David in a line. He will call you out for it. On 'Curb Your Enthusiasm' (Sun., 10PM ET on HBO) he berated a woman for doing a "chat and cut" in the line for the buffet table. She'd managed to push ahead of him by chatting to the guy in front of him and then sidling into the line behind the guy. Larry tapped her on the shoulder and congratulated her on her "excellent attempt at a 'chat and cut.'" In fact, she was so good at it that if Larry -- a self-professed master of the chat and cut -- hadn't been behind her, then she would probably have gotten away with it. http://www.aoltv.com/2011/08/08/larry-intervenes-in-a-chat-and-cut-on-curb-your-enthusiasm/
CURB YOUR ENTHUSIASM "Vow of Silence" Season 8 Episode 5 – Watch a preview and check out photos of the upcoming episode of CURB YOUR ENTHUSIASM which airs on Sunday August 7 at 9:00 PM ET/PT on HBO. Episode Synopsis: Curb Your Enthusiasm CURB YOUR ENTHUSIASM "Vow of Silence" Season 8Episode 5 – A failed alibi has coastal consequences; the Greenes' dog is denied a last meal; Larry (Larry David ) confronts a buffet-line transgressor. Read More... http://www.daemonstv.com/2011/08/06/curb-your-enthusiasm-vow-of-silence-season-8-episode-5/
Larry David had a dream on 'Curb Your Enthusiasm' (Sun., 10PM ET on HBO). He dreamed that he could date a woman at a restaurant he frequented, and then continue to go there after they broke up. His friends said it couldn't be done, but he was determined to do it. Then, as Larry does, he facilitated their break-up by putting expectations on her that weren't met. His argument about her giving back the $60 he lent her after she didn't use it, though, was a valid one. His mistake was in trusting her to put sun block on his head while this argument was happening. As such, she gifted him with the first thing he complained to her about. Smiley faces in text messages. It would have been that much funnier had she drawn it sideways on his head, to match the text version, but it was still a great visual ... particularly because Larry's next stop was the funeral of his assistant's father. http://www.aoltv.com/2011/08/01/larry-awkward-sunburn-funeral-curb-your-enthusiasm-video/
Watch a preview and check out photos of the upcoming episode of CURB YOUR ENTHUSIASM Season 8 Episode 4 titled "The Smiley Face" which airs on Sunday July 31 at 9:00 PM ET/PT on HBO Episode Synopsis: Curb Your Enthusiasm "The Smiley Face" Season 8 Episode 4 – Larry vows to topple a sacred dating taboo, and regrets making concessions to his new office neighbor; Jeff's alibi to get out of a dinner backfires. Directed by Jeff Schaffer. Read More... http://www.daemonstv.com/2011/07/31/curb-your-enthusiasm-the-smiley-face-season-8-episode-4/
Curb Your Enthusiasm keeps building. In its third week, the Larry David 's HBO comedy – which has always had a small but dedicated audience – was watched by 2.4 million viewers. That's a 24 percent jump compared to last week's episode for the most-watched installment of Curb in more than seven years (since March 2007). An additional 554,000 viewers tuned in for the 12:30 a.m. repeat, bringing the gross audience last Sunday to 2.9 million viewers. Read More... http://feedproxy.google.com/~r/thr/television/~3/IjghQ3mLWrk/curb-your-enthusiasm-has-watched-215506
In all honestly, we're going to have to back Larry David on the latest 'Curb Your Enthusiasm' (Sun., 10PM ET on HBO). Of course, social etiquette probably says that when the woman started screaming and reaching for the cake, you let her have the cake. But she did tell him to not let her have any, "no matter what." She'd just successfully lost 60 pounds on a diet, and it wasn't even a passing conversation when she made Larry her "dessert referee." There were several exchanges, leading him to believe she was serious. Maybe dessert is a particular weakness of hers, so Larry was more than happy to play the bad guy and not allow her to have a piece of cake. In fact, we'd argue that it does seem to be a weakness, and she has very little control over her impulses. Who fights that violently for a piece of cake? http://www.aoltv.com/2011/07/25/larry-dessert-referee-curb-your-enthusiasm-video/
Watch a sneak peek and check out a photo of the upcoming episode of CURB YOUR ENTHUSIASM Season 8 Episode 3 titled "Palestinian Chicken" which airs on Sunday July 24 at 9:00 PM ET/PT on HBO Episode Synopsis : Curb Your Enthusiasm "Palestinian Chicken" Season 8 Episode 3 – Larry plays "social assassin" to the hilt at a dinner party, on the golf course , and at a Palestinian restaurant with chicken to die for. Directed by Robert B. Weide. Read More... http://www.daemonstv.com/2011/07/24/curb-your-enthusiasm-palestinian-chicken-season-8-episode-3/
In its second week on the air, HBO's Curb Your Enthusiasm just keeps going up . For the second episode of the eighth season, the Larry David comedy improved from its opener last week, becoming the most-watched episode of the series in more than seven years. About 1.9 million viewers tuned in to the 10 p.m. telecast, up 12 percent from last week's premiere, and an additional 556,000 tuned in to the encore telecast at midnight. Cumulatively, Curb Your Enthusiasm lured a total of 2.5 million viewers, up 17 percent to last week. Read More... http://feedproxy.google.com/~r/thr/television/~3/zf0Yf0NB3lU/hbos-curb-your-enthusiasm-outperforms-213010
Sometimes, you just need your Chubby Hubby. And sometimes you have to go through someone who is sobbing uncontrollably in the freezer aisle to get it. That was Larry David's challenge Sunday on 'Curb Your Enthusiasm' (Sunday, 10PM on HBO), and he tackled it with typically awkward results. It's not like Larry didn't try to be understanding. He politely stated his intention, furrowed his brow and tried to look sympathetic. He even made little "aww" sounds. They refused to move. So he tried to go around them from the side to reach his Ben & Jerry's fix. The consoling friend wouldn't have it, and caught him mid-reach. "Sir, excuse me, now, I've tried to be nice," she said. "I have asked you to give us some space." http://www.aoltv.com/2011/07/18/crying-woman-keeps-larry-ice-cream-curb-your-enthusiasm-video/
Watch a sneak peek and check out photos of the upcoming episode of CURB YOUR ENTHUSIASM Season 8 Episode 2 titled "The Safe House " which airs on Sunday July 17 at 9:00 PM ET/PT on HBO Episode Synopsis: Curb Your Enthusiasm "The Safe House" Season 8 Episode 2 – Larry becomes an unlikely role model for battered women; Richard Lewis's relationship with a burlesque performer is put to the test; Leon survives a case of mistaken identity. Directed by Bryan Gordon. Read More... http://www.daemonstv.com/2011/07/15/curb-your-enthusiasm-the-safe-house-season-8-episode-2/


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Name: Illusions Of A Lady
This is a little seen classic from Jonas Middleton, who also directed the excellent Through the Looking Glass. Although no where near as ambitious as that film, this is an enjoyably kinky flick with the terrific Andrea True looking great and a memorably crazy scene from Jamie Gillis wearing women's underwear while jerking off and then covering on a pair of panties.
Actors: Jamie Gillis,Roger Caine,Davey Jones
Categories: 70s porn film, Helen Madigan, 1973, United States, English, Jonas Middleton, Helen Madigan, Andrea True, Michelle Magazine, Martine Gay, Jamie Gillis, Roger Caine, Davey Jones, Facial
Actress: Helen Madigan,Andrea True,Michelle Magazine,Martine Gay
Director: Jonas Middleton
Language: English
Duration: 58 min
Year: 1973
Country: United StatesHelen Madigan in 70s porn film
Sunday, September 25, 2016
Keywords:70s porn filmHelen Madigan1973United StatesEnglishJonas MiddletonAndrea TrueMichelle MagazineMartine GayJamie GillisRoger CaineDavey JonesFacial
© 2017 VINTAGESEXMOVIES.MOBI - Free Classic Porn Videos & Photos at VintageSexMovies.mobi : Watch and download all for free.


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No Matter How Beautiful A Car You Find, Not Buying It Just Because You're Short Of Enough Money, Can Leave You Heartbroken.
If The Vibration Changes Or Goes Away While Maintaining The Same Vehicle Speed, Then The Problem Is Not Related To Engine Rpm.
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From Complete Removal To Ensuring Dents Are Less Visible, Metal Levers Are Used To massage Out Dents From Behind Body Panels.
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Alpinetrek expert Christian
"This product is no longer available."
This is either an older model or we can or will no longer reorder this product from the manufacturer.
Don't give up – we have some other alternatives for you to choose from:
Rab - Women's Strata Hoodie - Synthetic jacket
Material information & features
- Suitable for:
- Design:
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- Material:
- Pertex Microlight (100% polyamide); DWR
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- Closure:
- full length front zip with chin protection
- Pockets:
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- Hood:
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- Weight:
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- Extras:
- elastic sleeve cuffs; Drawstring in the seam; Inside pocket serves as stuff sack
- Item No.:
- 104-0256


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Registration is now open for VFF 2016! Save the date!
And here is some exciting news for you about this year´s Visegrad Film Forum: The 5th Visegrad Film Forum will take place at the Faculty of Film and Television and in KC Dunaj in Bratislava from April 5-9, 2016. Come to enjoy five days full of screenings of student films, master classes with renowned experts, case studies of interesting film projects, discussions, and workshops – all connected with a very u ...Čtěte více ›


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Thursday, April 02, 2015
April Love - Favourite flowers
Roses, always roses (because they're pretty much the only flowers I'm not allergic to!)
This post is part of the April Love series, the brain-child of Susannah Conway. In Susannah's words:
The theme of the challenge, as you've probably guessed, is LOVE — love for ourselves, our lives, the world around us. LOVE expressed in gratitude and kindness. February might have Valentine but we've got April LOVE :)If you'd like to get in on the action, you can sign up here.
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My favourite cold remedy
I've been knocked flat by a cold this week. Sore throat, sniffles, stuffy head, temp..... feeling very sorry for myself! In fact, it...
Blogging for peace today! If you want to join in the blogblast for peace, please visit Mimi at Mimi Writes In the words made famous by John ...
Those of you who have me on facebook will know that one of my favourite bloggers, Susan (aka Whymommy) from Toddler Planet passed away rece...
How awesome must it be to be young, energetic and free to roam? ( Not that I would actually want to trade my home and family commitments....


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Sunshine Advanced Mix#4 growing mix
Sunshine Advanced Mix #4 is meticulously formulated to meet the needs of professionals, while being easy to use for beginners. Sunshine Advanced Mix #4 is made with Canadian Sphagnum peat moss, coarse perlite, coconut coir pith (coir), and a completely organic starter nutrient charge that contains gypsum, dolomitic limestone, and an organic wetting agent.Sunshine Advanced Mix #4 also contains a plentiful helping of multiple strains of mycorhizzae for improved nutrient and water uptake, and better resistance to stress and disease.
Premier Pro-mix BX
PRO-MIX 'BX' MYCORISE PRO is a "general purpose" peat-based growing medium with vermiculite and perlite, used for the cultivation of a wide variety of plant species. MYCORISE PRO ensures stronger, healthier plants due to more developed root systems and better fertilizer uptake.
Grodan Grow Cubes
Made of the same high-quality stonewool as the larger cubes and slabs, these miniature Grodan Grow-Cubes fit any container size or shape. Grow-Cubes offer easy irrigation, optimal root aeration and last throughout crop production. For any growing application, stonewool offers speed, ease of use and cleanliness
Bio Bizz Coco
OMRI-Listed Bio Bizz Coco Mix makes organic gardening easy. Its absorbent coir fibers make it great for hydroponics or as an amendment to soil, and it has a pH of between five and six
Improve the quality of your soil by using perlite
Hydroton Clay Pebbles
© 2011 Organic Grow Hut • Website Design by Lisa Day Burbaugh, NSBWebEffects.com


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o Project Execution Planning
o Level of Detail (LOD) Assistance
o Project based model estimating (quantities)
o Scheduling and 4D logistics planning


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A Leg That Fits - a webinar on Geomagic Freeform by designer and artist, Natasha Hope Simpson
(Pre-recorded webinar)
Natasha, a leg-amputee, is pursuing her dream of using 3D software and printing from 3D Systems to research how to deliver affordable custom prosthetics and prosthetic fairings to a mass audience. She does this using Geomagic Freeform and SLA 3D printing by 3D Systems.
In the video, you will hear about Natasha's story and see a live demonstration of Geomagic Freeform.
Please fill out the following form to access the recorded webinar.


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Our ES-985 Small Pedal Attachments will fit our Radio Flyer Trikes, Kettrikes and many other kid-sized trikes and riding toys.
Not for use on bicycles. The rider will not be able to put a foot down on the ground to keep from falling. Riders should always wear a helmet.
Home | About
Us | Terms & Conditions | Dealer
Inquiries | Email Us


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NEW ORLEANS — The New Orleans Saints no longer have to worry about an unbeaten season.
NEW ORLEANS — The New Orleans Saints no longer have to worry about an unbeaten season.
The Dallas Cowboys took care of that on Saturday night.
Dallas sliced through the Saints defense on its opening two possessions to build a 14-0 lead, and then took advantage of uncharacteristically poor play by the Saints offense en route to a 24-17 win at the Louisiana Superdome.
Now that the Saints (13-1 overall) no longer have an unbeaten season to worry about, their focus is on bouncing back from their first loss of the season. New Orleans hosts Tampa Bay Sunday in the Superdome.
"Dallas did a good job and played a good game. There were things we didn't do well and have to do better especially me. I have to do a better job," Saints coach Sean Payton said. "It was a tough loss for us. With the way the schedule shakes out, we have an extra day (to prepare for next week), and we will pay attention to the film and come back the best we can."
When asked if he was relieved of not having to worry about an unbeaten season Payton said, "It's a disappointment when you lose. It is no type of relief."
Despite Saturday's loss, securing the NFC's No. 1 seed is within reach for the Saints. With Minnesota loss on Sunday night, the Saints need to beat either Tampa Bay or Carolina to secure the top seed.
While Dallas (9-5) was having its way with the Saints in the first quarter, New Orleans' offense played its worst half of football all season.
Dallas' defensive front constantly harassed Saints quarterback Drew Brees into mistakes as he threw an interception and later fumbled on a sack by Cowboys defensive end DeMarcus Ware. Dallas recovered the fumble and added a field goal for a 17-3 lead with 4 seconds left in the second quarter.
Brees passed for a minus-1 yard in the first quarter and 127 yards in the first half. For the game, Brees was 29-of-45 for 298 yards with one interception and was sacked four times. Also, New Orleans was just 1-of-7 on third-down attempts, while the Cowboys finished 8-of-15.
"Early on, we felt we couldn't get in a rhythm, so we are down 14-0 and ran six plays offensively," Brees said. "They've got a good defense and deserve a lot of credit. For us, things weren't clicking and they got some pressure, and we didn't handle it very well. We felt their edge rushers were very good, and we contained them early. But late in the game, they were able to pin their ears back.
"We need to get back to starting fast and get that tempo going. (Missing out on the unbeaten season) is a disappointment and you don't know how many chances you get to do that. It is tough. We fought so hard to get where we are and won games in so many ways. We wanted to make it happen for the organization, the fans and the city. But the fact is we are 13-1. We still have that first seed in the NFC in sight. We want to be playing our best football going into the playoffs."
While the Saints offense struggled with consistency, Dallas found little resistance from the Saints defense at times. Dallas finished with 439 total yards compared to 336 total yards by the Saints. Dallas also won the turnover battle with two fumble recoveries and an interception, while its offense committed none.
Trailing 24-3, the Saints rallied in the fourth quarter to cut it to 24-17 after a Mike Bell 1-yard run and a 7-yard scoring pass from Brees to wide receiver Lance Moore with 8:06 left to play.
Dallas seemed to regain the momentum as the Cowboys put together a seven-play drive deep into Saints territory and lined up for a field goal.
However, the Saints got a little luck thanks to the misfortunes of Dallas' field goal kicker. Nick Folk's 24-yard field goal attempt hit the left upright leaving it a 24-17 game and 2:16 on the clock.
Unlike the Washington game when a missed field led to a Saints overtime victory, there was no miraculous comeback this time. New Orleans' perfect season ended when Brees fumbled while being sacked by Ware and Cowboys defensive tackle Jay Ratliff recovered the ball at the Cowboys 48 with six seconds remaining.
"We talked about what is still in front us (home-field advantage). This will sting a little bit, but we have to get back to work next week," Payton said. "We've got to correct the mistakes the best we can. We struggled on third downs, and when you do that, you are going to have less snaps, and you are off the field. That is one area that stood out."


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Even with well-known sampling biases, the fossil record is key to understanding macro-evolutionary patterns. During the Miocene to Pleistocene in the Caribbean Sea, the fossil record of scleractinian corals shows a remarkable period of rapid diversification followed by massive extinction. Here we combine a time-calibrated molecular phylogeny based on three nuclear introns with an updated fossil stratigraphy to examine patterns of radiation and extinction in Caribbean corals within the traditional family Faviidae.
Concatenated phylogenetic analysis showed most species of Caribbean faviids were monophyletic, with the exception of two Manicina species. The time-calibrated tree revealed the stem group originated around the closure of the Tethys Sea (17.0Ma), while the genus Manicina diversified during the Late Miocene (8.20Ma), when increased sedimentation and productivity may have favored free-living, heterotrophic species. Reef and shallow water specialists, represented by Diploria and Favia, originate at the beginning of the Pliocene (5 – 6Ma) as the Isthmus of Panama shoaled and regional productivity declined.
Later origination of the stem group than predicted from the fossil record corroborates the hypothesis of morphological convergence in Diploria and Favia genera. Our data support the rapid evolution of morphological and life-history traits among faviid corals that can be linked to Mio-Pliocene environmental changes.
Explaining rapid diversification and speciation remains a central challenge to evolutionary biology [1,2]. Much work has focused on either understanding the ecology and phylogenetic history of species-rich systems that have recently diversified along ecological axes (e.g. adaptive radiations) [3], or looking for patterns of species change in the fossil record [4-8]. Taking the molecular phylogenetic/ecological approach alone, however, excludes information about extinct lineages that may substantially bias our ability to identify cases of rapid diversification [9]. Conversely, relying on the fossil record alone limits our ability to detect evolutionary relationships between fossil taxa and some shifts in ecological function that may not be apparent from fossil character states. Ultimately, a more complete understanding of the processes that drive rapid diversification will require historical information from both molecular and fossil data. By examining systems that show recent speciation within monophyletic groups, ecological differentiation, and a strong fossil record, we can begin to link past to present processes in the understanding of the evolution of diversity.
The marine Caribbean fauna provides rare examples of diversification of monophyletic lineages within the context of well-understood changes in biogeography, oceanography, and climate. The isolation of Caribbean populations from their Indo-Pacific counterparts started ~15-17Ma when the closure of the Tethys Sea cut off connections between the Mediterranean and Indo-Pacific [10]. Isolation was complete ~3.45 - 4.25Ma when the rise of the Isthmus of Panama severed all Caribbean connections to the Indo-Pacific [11]. The period leading up to closure of the isthmus during the late Miocene to late Pliocene was characterized by changing global oceanographic circulation patterns, leading to drastic environmental, ecological, and taxonomic shifts within the Caribbean basin. Not only did the cessation of gene flow between the Pacific and Atlantic Oceans lead to widespread vicariant speciation across the newly formed isthmus [12-14], but on the Caribbean side, the accompanying geological and oceanographic changes caused an overall decrease in depth, primary productivity and turbidity and an increase in salinity, temperature, and local environmental heterogeneity [11,15]. Fossil records of many marine taxa during this period show elevated levels of taxonomic turnover [11,16-21], suggesting that climatic and geological variables drove elevated rates of cladogenesis and extinction.
This taxonomic turnover is particularly striking in corals of the family Faviidae, where an examination of the stratigraphic ranges shows that all extant species originated nearly simultaneously during the Mio-Pliocene [22]. Moreover, for faviids, this recent radiation has resulted in impressive diversification of ecological and life-history traits [23,24]. Modern species of Manicina are representative of a free-living lifestyle adapted to sediment-rich seagrass habitats that expanded during the Miocene then contracted during the Plio-Pleistocene [15]. In contrast, species of the brain coral genus Diploria tend to be reef-builders, dominating shallow water reef platforms in Pleistocene and modern times [25-28]. These two "sediment" and "reef" clades appear to share a common ancestor and ecological diversification seems to have occurred over a short period of geological time, suggesting it is tied to the contemporaneous increase in environmental heterogeneity [29]. Yet this punctuated diversification event is inferred from a fossil record, which may be incomplete or contain uncertainties in dating and taxonomic relationships that may influence our interpretation of past patterns.
Molecular data combined with well sampled fossil records provide opportunities to test existing evolutionary hypotheses and extend our understanding of both the tempo and mode of evolutionary diversification. In the Scleractinia, deep divergences between coral orders, suborders and families are increasingly well understood [30-33]. Yet a recent series of phylogenies exploring relationships at the familial level and below have demonstrated pervasive polyphyly and paraphyly at the generic level [34-39]. In addition, these studies have shown that between ocean basins, species group geographically rather than taxonomically [35,38,39]. In particular, Atlantic lineages of Faviidae and Mussidae appear to be more closely related to other Atlantic lineages than to congeners or even confamilials in other ocean basins. This geographic split supports the evidence from the fossil record of a radiation of the Caribbean coral fauna before complete isolation from the Pacific. However, the failure to resolve species relationships within the traditional coral family Faviidae, and a long history of taxonomic difficulties in identifying and classifying corals [32,36,40] demands an independent assessment of trends apparent in the fossil record.
To explore the tempo and mode of this evolutionary diversification, we unite a new multi-locus phylogeny of the Caribbean Faviidae with new stratigraphic compilations from the fossil record. Our well-sampled phylogeny allows Bayesian approaches to place these relationships into a temporal context by dating divergence times based on molecular data and fossil calibrations. We compare our time-calibrated phylogeny to temporal patterns of origination and extinction revealed by the Neogene fossil record, and find remarkable congruence between data sets. The timing of events revealed by this analysis strongly implicates paleoenvironmental changes as drivers of diversification in scleractinian corals.
We sequenced three single copy nuclear loci for six ingroup and one outgroup Caribbean faviid species. A total of 48 unique alleles were identified for CaM (alignment length=507bp), 38 alleles were identified for MaSC-1 (alignment length=490bp), and 55 alleles were identified for Pax-C (alignment length=418bp) (Additional file 1). Maximum likelihood and Bayesian analysis of gene trees showed little support for structure above the species level with no conflict between trees at highly supported nodes (Additional file 2). The taxa Manicina areolata and M. mayori shared some alleles at all loci, and unique alleles isolated from Diploria clivosa and D. strigosa did not always form monophyletic groups. A total of 94 individuals with unique genotypes were successfully sequenced at all three loci and used for a concatenated phylogenetic analysis. See Additional file 3 for genotype data of all individuals in study.
Bayesian and maximum likelihood trees had identical topologies at all major nodes with support values (Bayesian/ML bootstrap) indicated in Figure Figure1.1. The ingroup node was well supported (100/100) as well as species nodes for C. natans (100/100), D. clivosa (100/100), D. labyrinthiformis (100/98), D. strigosa (100/98) and F. fragum (100/100). Manicina mayori and Manicina areolata failed to form monophyletic groups, though support was high at the genus node for Manicina (100/94). The genus Diploria failed to form a monophyletic group. Diploria clivosa formed a clade with Manicina spp. and D. strigosa formed a clade with Favia fragum. Support for these nodes, however, was low (72/68 and 76/60 respectively).
BEAST analysis of the data produced a tree topologically consistent with those of the MrBayes and RaxML analyses. Visual inspection of plots in Tracer v1.5 [41] showed rapid convergence of the analysis and narrowing of priors with all parameters having an effective sample size (ESS) of >1900. The mean rate of substitution was 6.77× 10-4 per site (95% Highest Posterior Density (HPD) interval: 4.49 × 10-4 - 9.06× 10-4) with a coefficient of variation of 1.17 (95% HPD interval: 0.66- 1.72) indicating significant heterogeneity in substitution rates across the tree.
Mean ages of species, ingroup, and root nodes with 95% HPD intervals are shown in Figure Figure2A2A and listed in Table Table1.1. The posterior mean of the time of the most recent common ancestor (TMRCA) of the Manicina group, which was calibrated from species fossil data, shifted several MY from the prior distribution, indicating that the sequence data is influencing divergence dates. For D. clivosaD. strigosa, and F. fragum, mean estimated ages fell close to the earliest possible dates of their appearance in the fossil record. For the D. labyrinthiformis and Manicina nodes, fossil dates were closer to the youngest part of the 95% HPD interval. Mean origination time of D. labyrinthiformis is pushed back approximately 1.6 MY earlier than previously seen in the fossil record, putting it closer to the origination times of the other species. All mean species origination dates occur shortly prior to the final closure of the Central American Isthmus at 4.25 - 3.45Ma [11], but we note that the youngest part of 95% HPD for F. fragum and D. labyrinthiformis overlap with this estimated age of final closure. The timing of the Manicina node is considerably earlier than the appearance of the Manicina areolata in the fossil record, indicating that this genus diverged earlier than the first appearance of M. areolata. Deeper nodes in the tree had significantly larger HPD confidence intervals, due to the lack of fossil calibrations for earlier taxa. The estimate of origination time for the ingroup was 14.10Ma (95% HPD interval: 8.77-20.09), while origination of the entire Caribbean Faviidae group is indicated by the root node at 17.56Ma (95% HPD interval: 10.04-26.44). These dates coincide with the timing of the closure of the Tethys Sea in the eastern Mediterranean [10].
Overlay of the molecular phylogeny onto the fossil stratigraphy reveals three striking patterns (Figure (Figure2B).2B). First, older and extinct Diploria and Favia cannot be reconciled with this molecular tree, suggesting these genera are not monophyletic. Second, the origination and diversification of a clade of sediment dwelling corals (particularly Thysanus and Manicina) is confirmed by both the fossil record and molecular phylogeny. Lastly, the appearance of new reef dwelling species of Favia and Diploria is simultaneous in the fossil record around 5Ma.
Thorough sampling of individuals within species in our combined phylogenetic analysis confirms that most modern Caribbean species form well-supported monophyletic lineages (Figure (Figure1).1). This allows us to reject the idea that widespread hybridization on ecological time scales [42] is important to the evolution of Caribbean faviids, though limited introgression not detected by this data set might have played a creative role in adaptive processes [43]. The exception lies within the two modern species of ManicinaM. areolata and M. mayori, where extensive allele sharing between species might indicate sub-species status. While M. areolata is a seagrass specialist, and drifts remarkable distances on the sediment surface as a free-living adult [44], M. mayori is a rare reef species that remains permanently attached as an adult. In Panama, these two morphologically distinct species co-occur within sites yet segregate ecologically by depth-related habitats. An approach that combines ecological/reproductive comparisons, morphometric data, and further genetic analyses such as the coalescent-based model of isolation and migration [45] could resolve this issue.
Above the species level, we could not further resolve the branching order of species within the larger clade. Previous single locus phylogenies using mitochondrial and nuclear genes that have included this group have shown a similar lack of resolution within the Caribbean faviids[35,36]. Another study by Nunes et al.[39] shows some supported structure within this group. However, as this paper was looking mainly at broader scale phylogeographic relationships, sampling was done on only few individuals per species within the Caribbean faviids and using only a single mitochondrial marker and a single nuclear marker. For examining relationships below the familial level, the low rates of mtDNA evolution in corals might limit the ability to detect more complex topologies amongst these species. With the increased sampling sizes of multiple loci with higher levels of variation (Additional file 2), we found little evidence for monophyly within genera, and branch lengths tended to be long (Figure (Figure1).1). Therefore, our inability to resolve relationships among species is consistent with rapid diversification and short internal branch lengths deeper in the tree. With the rapidly declining cost of high throughput sequencing, a phylogenomic approach [46,47] for this set of taxa is likely to improve topological resolution.
Molecular divergence dating indicates extant Caribbean "faviid" corals radiated rapidly during the late Miocene to early Pliocene (Figure (Figure2).2). This ecological radiation coincides with a series of biological and physical changes in the structure of shallow marine habitats during the early geological development of the Isthmus of Panama. During the Late Miocene, shallow marine habitats were dominated by broader and more gently sloping sedimentary shelves [48], while productivity in the water column above was much higher compared to the modern productivity of the Caribbean Sea [49]. Klaus et al.[15] hypothesize that these extensive mesophotic sedimentary bottoms may have selected for free-living coral species with large tentacle morphologies that were efficient at heterotrophic feeding. Interestingly, our node age for the clade containing the two living Manicina species is 8.21Ma, which coincides with the appearance of other sibling Manicina species in the fossil record that have since gone extinct [22]. Thus, it appears we are sampling the evolutionary remnants of a once more diverse and ecologically dominant clade. As the Miocene transitions into the Pliocene, the increasingly isolated Caribbean Sea becomes more oligotrophic and the once broad shelf habitats are now dominated by steeper reef platforms, ideal conditions for primarily photoautotrophic reef species. Our time-calibrated phylogeny shows repeated speciation events of Diploria and Favia species between~4 – 6Ma that are either reef specialists or are limited to very shallow (< 5m) seagrass habitats. Thus the fossil record and molecular data broadly agree on the timing of these ecological radiations, which are temporally correlated with changes in habitat structure and productivity.
Deeper in the tree, node ages for the stem groups of the Caribbean faviids correspond to the isolation from the Mediterranean during the closure of the Tethys Sea (Figure (Figure2A).2A). While these dates support the widely accepted notion of divergence driven by increased isolation of the region, the radiation of the stem group is much later than indicated by the fossil record (Figure (Figure2B).2B). The origination of the FaviaDiploriaManicina (FDM) clade is in the early Miocene, but older Oligocene Diploria fossils and Eocene Favia fossils are more distantly related, suggesting that both genera are para- or polyphyletic. Ken Johnson reached a similar conclusion based on morphological differences [22], hypothesizing that early Diploria and Favia are unrelated to their modern morphological counterparts. Morphological convergence appears to be a common theme in coral evolution [35] and our analysis points out some of the difficulties in determining the systematic positions of extinct taxa. The use of more informative micro-structural characters that can be quantified in both living and fossil species may be a promising approach to this problem [30].
Congruence of morphology, stratigraphy, and estimates of node ages can be used to include fossil taxa into potentially monophyletic lineages. For example, the diverse members of living and fossil taxa of the genus Manicina form a well-supported monophyletic group in Johnson's morphological phylogeny with all fossil origination dates falling within the lower confidence interval for the molecular Manicina node age (Figure (Figure2A).2A). Superimposing the age-calibrated molecular phylogeny onto stratigraphy significantly alters the interpretation of the speed of evolution in this group (Figure (Figure2B),2B), indicating rapid diversification of sediment dwelling corals in the late Miocene.
Our time calibrated phylogeny confirms fossil evidence that extant Caribbean coral species originated during a period of lineage diversification between 4 and 6Ma (Figure (Figure2).2). This diversification event corresponds with ecological radiation into three main ecological niches exemplified by modern Caribbean faviids [21]: (i) small, free living morphologies adapted to sedimentary environments (ii) attached species that live in shallow rubble beds and patch reefs, and (iii) massive colonies the build forereef slopes (Figure (Figure1).1). During the same period, we also see diversification of reproductive strategies [23], from tightly synchronized annual mass-spawning events and broadcasting larvae typical of Diploria[24] to multiple lunar cycles of reproduction and brooding development found in Favia and Manicina[50,51].
The changes in morphology and life history coupled with widespread environmental changes are suggestive that diversification of Atlantic "faviid" coral might be driven by the evolution of adaptive traits. Using our current phylogeny as a stepping stone, increased genomic and taxonomic sampling of Atlantic corals should allow us to take advantage of several promising new approaches to estimate rates of diversification and evaluate models of adaptive radiation [52,53].
By combining data from the fossil record with molecular phylogenetic techniques for the first time, this study has given us extensive insight into the tempo of diversification in an ecologically diverse group of Caribbean corals. Two separate lines of evidence now verify the existence of a Mio-Pliocene radiation, while we have been able to additionally confirm species identity, verify origination dates, and understand taxonomic relationships in this diverse and ecologically important group. These findings give us the tools to re-interpret trends seen in the fossil record, allowing us to begin to link patterns of macroevolution to paleoenvironmental changes and gain a new comprehension into the origins and drivers of diversity in the Caribbean.
Besides clarifying evolutionary history, this study has broader contemporary implications. With global change currently causing a rapid decline in coral reef populations around the world [54,55], understanding the processes that generated diversity in coral species will be key to predicting future changes and directing conservation efforts [56]. It has been suggested that species that evolved in a more heterogeneous environment and survived past climatic fluctuations will be more resistant to current global change [29]. Understanding patterns of Caribbean coral evolution during the Pleistocene may be key to understanding the potential outcomes of current environmental impacts.
We sampled six of the seven nominal species from the genera FaviaDiploria, and Manicina that form a monophyletic group within the Caribbean Faviidae [35,39]. The single missing taxon is Favia gravida, closely related to Favia fragum, but with a distinct non-Caribbean distribution in that it has been only described from Brazil and West Africa [57,58]. We used the genus Colpophyllia as the outgroup because it has previously been shown to be a stem taxon to the ingroup species [35,39]. Extensive sampling within each species was conducted at two reef systems in the Caribbean Sea: the Bocas del Toro, Panama, and the Florida Keys, USA with additional F. fragum sampled from St. Croix, USVI. The complete list of samples and collection localities is provided in Additional file 3. Skeletal vouchers are deposited in the University of Iowa Paleontology Repository (http://geoscience.clas.uiowa.edu/paleo/index). Samples were collected, preserved, and genomic DNA extracted as described in Carlon and Lippé [59]. Skeletal vouchers were processed by bleaching in a 50% hypochlorite/water solution overnight, rinsing in DI water, and thoroughly drying. Species identification was conducted by D. Carlon in the field and confirmed by A. Budd from vouchers. Complete descriptions of these taxa, photos, and references are available from the Neogene Marine Biota of Tropical American (NMITA) database (http://eusmilia.geology.uiowa.edu/).
For this study we chose to focus on nuclear markers, since rates of mitochondrial DNA evolution have been shown to be very slow in corals, limiting the ability to detect more recent speciation events [60]. We amplified three single-copy nuclear regions with primers listed in Additional file 4. Pax-C and CaM primers target introns located within the Pax protein and calmodulin binding protein respectively[61,62], while MaSC-1 is an anonymous region originally sequenced in Montastraea annularis[63]. For PCR amplification of all three loci, we combined: 1μl of 1x to 100x diluted genomic DNA with a 24μl PCR master mix consisting of: 0.3μl of each primer (10μM), 1μl dNTPs (2.5mM each), 2.5μl 10x reaction buffer, 1μl MgCl2 (25mM), 1μl BSA (10mg/ml), 0.3μl Taq polymerase (Bioline), and 17.6μl of H20. Each reaction was run at 95°C for 10min, 30 cycles of 94°C for 30s, Ta for 40s and 72°C for 1min, with a final extension of 72°C for 10 minutes. Purified PCR products were sequenced on ABI 3731 XL 96 capillary DNA analyzers at the University of Hawaii at Manoa and chromatograms were then analyzed and edited using Sequencher 4.5 (Gene Codes). Direct reads revealed indels segregating within many of the species, and preliminary cloning verified multiple indels within all three genes. Since phasing length-variant heterozygotes (LVHs) from direct reads proved unreliable, we cloned 86% of the PCR products from individuals with LVH phenotypes. We cloned PCR products using a TOPO TA cloning kit (Invitrogen) and sequenced using standard M13 primers. Single nucleotide polymorphism heterozygotes were phased using the software PHASE v2.1.1 [64,65]. We used the non-recombination model, and phase thresholds of 0.90. To convert data between FASTA and PHASE formats, we used the webtool SeqPHASE [66]. Haplotype sequence data are available as Genbank PopSets. Accession numbers are listed by species in Additional file 1.
Allele sequences were aligned automatically using MAFFT v6 [67,68], and corrected by eye in MacCladev4.08 [69]. Indels were coded as missing data. Models for molecular evolution for Bayesian analysis for both gene trees and the partitioned tree were selected using the Akaike Information Criteria (AIC) in jModelTest v0.1.1 [70,71]. For the gene trees, the best fitting model for the CaM and Pax-C alignment was GTR+G, and for MaSC-1 the model was GTR. Bayesian trees for all three loci were generated in MrBayes v3.1 (5,000,000 generations, nruns=2, nchains=4) [72,73]. Trees were sampled every 100 generations and 5,000 trees were discarded as burn-in. Maximum likelihood analysis was performed using RaxML 7.2.6 [74,75] with 1000 rapid bootstraps using the default GTR+G model for all loci at the recommendation of the programmers.
Individuals sequenced at all three loci were used for the construction of a combined partitioned tree. For heterozygotes, SNPs were coded as ambiguous data using standard IUPAC nucleotide ambiguity codes. For the Bayesian analysis, the K80+G model was chosen for the CaM partition, the HKY+I+G model for theMaSC-1 partitions, and the HKY model for the Pax-C partition. Bayesian trees were generated using MrBayes v3.1 (20,000,000 generations, nruns=4, nchains=4). Trees were sampled every 1000 generations and 5000 trees were discarded as burn-in. Maximum likelihood analysis was performed using RaxML v7.2.6 with 1000 rapid bootstraps on the Cipres Web Server [76]. For this analysis, the default GTR+G model was used as above.
The program BEAST v1.7.1 [77] was used to estimate divergence dates at species nodes using available fossil data for calibration. Input files for the analysis were generated with Beauti v1.5.4 using a partitioned alignment file of 80 individuals. We used a Yule process speciation prior for branching rates along with an uncorrelated lognormal model for a relaxed molecular clock. Models for molecular evolution for BEAST analysis were selected using the Akaike Information Criteria (AIC) in MrModelTest v2.2 [78]. The HKY+G model was used for the CaM and MaSC-1 partitions, and the HKY+I model for the Pax-C partition. Base frequencies were estimated throughout the analysis. Based on the phylogenetic analysis, all species nodes were constrained to be monophyletic except for M. areolata and M. mayori, which were constrained at the genus node. Shape parameter priors were taken from MrModeltest v2.2 and priors for rates of evolution and Yule birth rates were chosen based on defaults narrowed from preliminary runs.
Stratigraphic ranges of extinct and living Caribbean Faviidae were compiled from the literature and unpublished sources (Additional file 5). Fossil stratigraphic ranges for extant species were used to calibrate species nodes for Diploria spp., Favia fragum and Colpophyllia natans and the genus node for Manicina (Table (Table2).2). Calibrations of nodes were done following the guidelines of Ho and Phillips [79]. For all date priors, we used a lognormal distribution with a hard minimum bound set at youngest possible date of first appearance in the fossil record. The mode of the distribution was set to be slightly older than the oldest possible date of first appearance. Finally, the 95% probability distribution was set to encompass a soft maximum bound at the time of the closure of the Tethys (~17Ma). These distributions incorporate the best-known estimates for origination dates of these taxa, but are wide enough to allow for shifts in dates that may reflect errors due to interpretation or incompleteness of the fossil record.
BEAST was run 4 times (generations=20,000,000, samplefreq=1000) on the Bioportal webserver at the University of Oslo [80]. Log files were examined in Tracer v.1.5 [41] to assess convergence of each run. After a 10% burn-in was removed, logs and trees for all runs were combined in LogCombiner v1.7.1 and chronograms were generated with Tree Annotater v1.7.1.
The authors declare they have no competing interests.
SAS designed the study, generated molecular sequence data, ran phylogenetic analyses, and drafted the manuscript. AFB confirmed morphological identification of species, compiled stratigraphic data, and helped draft the manuscript. DBC participated in the study design, collected field samples, ran phylogenetic and phasing analyses, and helped draft the manuscript. All authors read and approved the final manuscript.
Alleles and Accession Numbers by Species. Number of individuals sequenced per species (n), the number of alleles isolated per locus per species, and Genbank accession numbers. All species carried unique alleles, except for the two Manicina species. The two last rows give the number of unique alleles in the combined Manicina data sets (Manicina spp.) and the combined 6 ingroup species. (PDF 42 kb)
Gene trees for (A)CaM, (B)MaSC-1, and (C)Pax-C. Alleles are designated by locus_allele number. Node labels indicate Bayesian posterior probabilities/Maximum likelihood bootstrap support, --<50% ML support. Two-tone boxes indicate alleles shared between taxa. All trees produced in MrBayes v3.1 (generations=5,000,000, nruns=2, nchains=4.) The models of evolution were GTR+G for Cam and Pax-C and GTR for MaSC-1. See Additional File 3 for individual genotypes.
Sampling and genotype data for all individual corals. Samples and multilocus genotypes used in gene and concatenated trees. Heterozygous genotypes that could not be resolved by cloning or PHASE 2.1.1 are indicated as '?/?+'; and '0' indicates PCR failure or poor sequence quality. The two last columns designate which samples were used in the concatenated ML/Bayes trees (Figure (Figure1)1) and the BEAST analysis (Figure (Figure22).
Primers used for direct sequencing. Sequences and annealing temperatures for primers used in this study.
Stratigraphic Ranges of the Fossil Caribbean Faviidae. Compiled first and last occurrence data, references, and notes for all Caribbean fossil faviid species.
We thank S. Barnes, J. Coloma, J. Fitzpatrick, and S. Walls for help collecting in the field. Brendan Holland kindly provided Diploria samples from the Flower Garden Banks. Collections from St. Croix were funded by a grant from the University Research Expeditions Program, University of California (UREP) and we thank all expedition participants for excellent field assistance. Allen Chen's lab graciously provided the original Calmodulin primers. Thanks to Joanna Bince for her assistance in sequencing and cloning introns, and George Roderick for lab space and computer time for the final portions of the analysis. All corals were collected by permits from the Republic of Panama (ANAM and CITES) and the Florida Keys National Marine Sanctuary. Our field work in Panama benefited from the extremely helpful staff of the Smithsonian Tropical Research Institutes Bocas del Toro Station. This research was supported by a grant from the National Science Foundation (DEB-05-43661) awarded to DBC.
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