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Results tagged ' Gary Matthews Jr. '
Matthews seeking new home
As he was packing his bag at Yankee Stadium late Sunday night, a long season over, Gary Matthews Jr. looked up and said, "It's time."
It was not the first time he'd said this, and his meaning was clear. It's time, in Matthews' mind, to move on. Time to find a place where he can play center field every day and take full advantage of his multiple skills at age 35.
Tired of being a fifth wheel in the Angels' outfield, Matthews desperately wants to wear a new uniform with two years remaining on a five-year deal that brought him $50 million after his 2006 All-Star season in Texas.
He thinks there are teams that can see what he has to offer, and he hopes something can be done to make it happen. The $23 million owed him complicates the situation, but there are possible fits with clubs that have high-priced contracts they could move in exchange for a versatile switch-hitter who can play high-caliber center field.
Matthews had one superb half in '07, leading off and batting cleanup and making all the plays in center, before a knee injury set him back and forced him to end his season in civvies while the Angels were getting swept by Boston in the ALDS.
They went out and acquired Torii Hunter that winter, a move not even Matthews could criticize. A smart baseball guy, having grown up in the game with a slugging father, Gary Sr., he understands Hunter's tremendous value on and off the field. It just happened that Torii is one of the elite players in the game at Matthews' natural position.
Life as a backup role player has been unfulfilling for the son of Sarge.
Matthews had a nice little run this season when Hunter was sidelined for a month with a groin injury sustained banging into outfield walls. During that time, Matthews was renewed, emotionally and physically, and it showed in his performance.
Carrying the momentum of a strong June finish, the Angels were 17-9 in July with Hunter missing all but seven games. During one memorable stretch with Hunter and Vladimir Guerrero both sidelined, the Angels were 17-3.
The Angels were 32-13 with Matthews starting in center field, and he batted a robust .358 with runners in scoring position, when his competitive juices were flowing.
Those numbers might be a more accurate reflection of what he can do than his pedestrian .250 batting average, .361 slugging and .336 on-base numbers for the full season, with an irregular work load.
It took him a while to find his stroke after Hunter went on the shelf, but once he did, Matthews carried it to the finish with strong performances in August (.290 average, .452 slugging, .389 on-base percentage) and September/October (.286/.457/.444, respectively).
His play in center field is far superior to what he does in left and right, where he struggles at times with the hooks, slices and angles. In center, Matthews ranks among the top third in the game in the studied view of Hunter, the master.
Teams with needs in center should seriously explore taking on a player who has plenty of game left and a hunger to show what he can do.
This certainly isn't anywhere near the top of the Angels' winter agenda, with seven free agent cases to weigh along with eight arbitration-eligible players to satisfy. But at some point, dealing with Matthews – and dealing him in a mutually satisfying manner – would seem to be the right thing to do.
Guerrero gets a day off
Vladimir Guerrero, like fellow cleanup man Alex Rodriguez from the other side, was given a day off on Wednesday as the Angels and Yankees got together in the bright sunlight of Angels Stadium for their final regular-season date.
They could meet again in October, and if that happens, neither Guerrero nor A-Rod will be skipping any at-bats or innings. With a scheduled day off on Thursday, Guerrero gets two days off his feet, which ought to restore some life in his legs.
"Just a day [off]," Angels manager Mike Scioscia said of his designated hitter, who is batting .299 (with 15 homers and 47 RBIs in 91 games) in his quest to reach .300 for a 13th consecutive season. "He'll be ready on Friday [against Oakland].
"He's played a long time with a lot of swings. You can talk about DHing, and obviously it's not as demanding as playing in the field. But there's still a lot that goes into what a guy has to do. Especially since he's been dealing with leg issues, which put him on the DL the second time."
Bobby Abreu was in the DH spot against A.J. Burnett, with Gary Matthews Jr. getting a start in right field.
Guerrero, who had right knee surgery last October, opened the season with a torn pectoral muscle that cost him 35 games when it was diagnosed on April 18. He returned on May 25 and sustained a strained lower right hamstring, in the upper calf area, making a play in right field, putting him back on the DL from July 8 to Aug. 4.
A total of 56 lost games cost him a realistic shot at his 12th consecutive season with at least 25 homers and a .300-plus batting average. He's with Lou Gehrig as the only players ever to do that 11 years in succession.
In the past 50 years, only Tony Gwynn and Rod Carew, spray hitters with speed, managed to bat at least .300 for at least 12 seasons in a row, a streak Guerrero started in 1997 on Montreal's carpet.
One more historical reference point underscores how truly unique this man has been in his amazing career. With a .322 career average and 407 home runs, Guerrero is in a club of six with Jimmie Foxx, Gehrig, Stan Musial, Babe Ruth and Ted Williams at .320-plus and 400-plus.
Someday, Vlad will join the legends in Cooperstown, N.Y., home of immortals.
Big night for Evans, Willits, Wilson
In the ninth inning of Thursday night's 4-3 win in Boston, as significant a victory as the Angels have produced this season in some respects, three September call-ups were on the field in support of Kevin Jepsen and Brian Fuentes – including the man calling the pitches, Bobby Wilson.
Reggie Willits was in left field, where Juan Rivera had opened the game, and Terry Evans was in right, which had been occupied by Gary Matthews Jr.
Willits dropped a perfect bunt to set up the winning run in the top of the ninth, scored by Evans as a pinch-runner, and Evans squeezed the final out.
Wilson blocked a few balls that could have been trouble and once again handled himself with confidence and poise at the most difficult position on the field.
"That's big for guys like us," said Evans, a graceful athletic with a lean but muscular frame, "to know they have the confidence to put us out there in a situation like that, a big game on the line. It gives us confidence, as well. It's huge."
Evans had another big year at Triple-A Salt Lake alongside Willits and Wilson, as well as Brandon Wood, Sean Rodriguez, Freddy Sandoval, Chris Pettit and the rest of the Bees' formidable lineup. Evans, Wilson and Wood are out of options, meaning they'll either be with the Angels next season or available to other clubs unless they're included in deals.
"Anything we can do to contribute, we're happy," Evans said. "It can be the smallest thing. For us, that's our role here. We have such a great lineup, we know what our roles are. And it's exciting to get a chance to make any kind of contribution."
Evans and Pettit have been used as pinch-runners late in games, freeing Willits for a role he is beginning to master: dropping a sacrifice bunt in conditions far more difficult than any casual fan would realize.
"It's something he's been great at, and it helps if they can hold him back for those spots," Evans said. "Chris can run, and Freddy can run a little bit too. Reggie can do so many things, he's a good guy to have around late in games. Plus, with Bobby catching, they can save [Mike] Napoli for pinch-hitting situations."
Wilson has caught 11 innings this season, his pitchers yielding two earned runs. He made a game-saving, ninth-inning save of a ball in the dirt in Oakland when John Lackey (nine innings) and Fuentes combined for a 1-0 shutout in 10 innings.
Wilson was sent to Salt Lake after that Aug. 4 game, making it a bittersweet day.
"I love it any time I put on the gear and get a chance to play," Wilson said. "We all want to be in there, and it was great that Terry, Reggie and I were all on the field together."
"Especially," a grinning Willits said, his bunt having set up Howard Kendrick's game-winning single, "when we win."
Evans had gone in to run for Rivera after his leadoff walk against Red Sox lefty Billy Wagner.
"With more bench strength, especially pinch-running," Angels manager Mike Scioscia said, "it's a way to infuse some speed into a situation. The baseball experience of the Major Leagues, cutting their baby teeth, is a big step for these younger players. September is important."
Roenicke defends Rivera's defense
The insiders and media sharks were swirling around Angels left fielder Juan Rivera on Thursday, taking him to task for not diving full-tilt for the pop fly by Alex Gonzalez that dropped in fair territory in shallow left field on Wednesday night at Fenway Park.
Gonzalez's bases-loaded single gave the Red Sox a controversy-riddled 9-8 win, coming after Nick Green had walked on a full-count, forcing home the tying run, after the Angels felt Brian Fuentes had him struck out twice, on a checked swing and again on a 3-2 fastball at the knees and over the middle of the plate.
Rivera didn't have much to say about it, other than, "I didn't think I could catch it." He went to a few teammates to see what they thought and found support.
Emphatic, unyielding affirmation came from outfield coach Ron Roenicke, manager Mike Scioscia's bench coach.
"I saw the replay," said Roenicke, a former Major League outfielder known for his defensive skills. "He's not going to catch that ball. So he pulled up.
"When you go for a ball and know you can't catch it in your mind, you pull up. I don't want them to dive for a ball if they know they can't get it and get hurt. From what I saw, he wasn't going to catch it."
Rivera didn't appear to be playing as shallow as center fielder Torii Hunter or right fielder Bobby Abreu, but the Green Monster can distort perspectives.
"We're playing shallow," Roenicke said. "We were in a little bit."
Scioscia was asked if he considered removing Rivera – who'd gone 3-for-5 with a two-run double putting the Angels in front in the seventh — for defensive purposes with the Angels leading by a run going into the bottom of the ninth. Reggie Willits and Gary Matthews Jr. would have been options.
"Vlad [Guerrero] was already out of the game [with a bruised rib cage after getting hit by a pitch], and Juan's swinging the bat well," Scioscia said. "Juan is comfortable out there.
"He went after it hard. If he thought he could have dived and caught it, he would. Didn't think he had a chance."
Rivera has had a solid season defensively in left, making several game-saving catches and unleashing strong, accurate throws with consistency.
Some outfielders, Roenicke said, are less comfortable diving full-tilt.
"Juan plays hard," Roenicke said. "If he thought he'd have caught it, he'd have come after it. I've never seen him prone dive. I've seen him slide into a wall for a catch. Torii will dive forward, but there are not many guys who will dive forward."
Roenicke said Rivera had not asked him about the play but anticipated that he would – and the coach's support would be forthcoming.
Hunter on comeback trail
Rancho Cucamonga's good fortune is Arkansas' bad luck.
Fans of the Angels' high Class A California League team get the treat the next few nights of watching one of baseball's premier players – Torii Hunter – work out the kinks after a month-long stay on the disabled list with an adductor strain on his right side.
Hunter thought he was ready to go play in his native Arkansas for the Angels' Double-A affiliate in Little Rock over the weekend, but that fell through when he didn't pass a strength test administered by the training staff in Chicago.
"That was really disappointing," Hunter said. "I wanted to play at home. But at least I'm getting close. I'm hoping to be back out there [in center field for the Angels] this weekend in Baltimore. I hate sitting around watching."
The Angels have held their ground in Hunter's absence with Gary Matthews Jr. doing a solid job in center field. Matthews got his first day off on Tuesday night, with Reggie Willits in center, since moving in on a regular basis on July 8.
Hunter is batting .305 with 17 homers and 65 RBIs. His slugging (.558) and on-base percentages (.380) would be career highs.
Hunter's injuries were the result of collisions with the walls at Dodger Stadium on May 23 and at AT&T Park in San Francisco on June 15.
A case can be made that Hunter was the MVP in the American League over the first half of the season, carrying his club offensively while playing the brand of defense that has made him a Rawlings Gold Glover winner eight years running. Angels color commentator and former Major League pitching star Mark Gubicza, one of the brightest talents in his new field, has made that observation.
Matthews feeling validated
The past month has brought validation to Gary Matthews Jr. With Torii Hunter sidelined, Matthews has started every game in center field during a 17-6 stretch by the Angels, contributing offensively and defensively.
"To be able to contribute to a winning team is something every player wants," Matthews said. "All any player can ask for is the opportunity. When you're able to go out and perform, help your team win, it feels good.
"It has reiterated the fact I can play every day and be successful. I've gotten some really big hits and been a key contributor to the team. It has reiterated what I've said and felt all along."
Coming into Thursday's series finale against the White Sox in Chicago, Matthews has hit safely in eight of the past nine games, batting .351 during that stretch with six runs scored and six RBIs.
His .247 batting average isn't impressive, but he has numbers that clearly demonstrate that he has elevated his game in clutch situations. Matthews is batting .295 with runners on base, .344 with runners in scoring position, .455 with two out and runners in scoring position, and .571 with the bases loaded.
"I've never been afraid of big situations," he said.
Matthews has talked with Chone Figgins about sharpening his focus with the bases clear, when his average slips to .206. But Gary also knows he excelled as a leadoff man in Texas in 2006, before signing his five-year free agent deal with the Angels, and that it's just a matter of getting back to that mindset.
"I don't check the numbers," Matthews said, "but as a player, you know what you're hitting in certain situations. I've been really comfortable hitting with runners in scoring position. I don't know if it's a matter of better concentration, but I've had some success in those situations that I'd like to carry over to all of my at-bats.
"Certainly it brings out your instincts in those big situations, but it's also the competitor in any player. There's something about having a runner on second base or third base. You know a pitcher steps it up a level, and as a hitter you have to step it up, too. In talking with Figgy about your approach with no runners on, it could be I'm trying to do too much in those situations."
As he continues his rehab from an adductor muscle strain in his right side, Hunter knows his job is in good hands.
"For a guy who didn't play regularly for three months," Hunter said, "Gary's playing at a high level. That's not easy, what he's done. He's in the upper class of defensive center fielders, and he's showing it. And he's been coming up with big hits.
"It's good you've got somebody like that who could be a starter anywhere else. We've got a lot of quality players on this team."
Sean Rodriguez impresses the boss
Perhaps no player better embodies the Angels' remarkable organizational depth and versatility than Sean Rodriguez, who is emerging as their new Chone Figgins with his ability to play capably all over the field.
"From the perspective of being an everyday player, he's opened up a new dimension being able to play the outfield every day," Angels manager Mike Scioscia said on Wednesday as Rodriguez was back with Triple-A Salt Lake following two superb games in left field in Minnesota over the weekend. "He's got enough thunder in his bat to play corner outfield. He can play second base every day, shortstop, third base. He's a baseball player, a terrific athlete.
"The way he handled himself on the defensive end [at the Metrodome] is a plus. We've seen glimpses of the power he has. He's putting it together."
The Angels elected to return Rodriguez to Salt Lake and keep Reggie Willits on the 25-man roster because of Willits' ability to do multiple things late in games — pinch run, get a bunt down, hit-and-run — and his superior experience in center field, where he'll support Gary Matthews Jr. in Torii Hunter's absence.
Rodriguez, a natural center fielder who played there at G. Holmes Braddock High School in Miami, transformed himself into a quality middle infielder after signing with the Angels in 2003 as a third-round choice in the First-Year Player Draft.
Developing his infield skills while hitting with power as he climbed the organizational ladder, Rodriguez returned to the outfield, playing all three positions along with the infield, to enhance his appeal as an all-purpose talent. It helped Figgins carve out his career, and now, having settled in full-time in one role, he's one of the game's elite third basemen.
Summoned to the big club from Triple-A Salt Lake, where he's been pounding Pacific Coast League pitching all season, all Rodriguez did in two starts in Minnesota was lash a pair of singles and make two excellent defensive plays in his first start in left field, then bang a homer over the center field wall the following day.
Arriving in Chicago, Rodriguez learned that he was heading back to Salt Lake to make room for Vladimir Guerrero, coming off the disabled list after recovering from a muscle strain behind his left knee.
"It was fun while it lasted," Rodriguez said. He couldn't hide his disappointment, just as Brandon Wood and Bobby Wilson days before him.
Scioscia raved about the impression made by the 24-year-old Rodriguez last season at second base during a lengthy stretch in May with Howard Kendrick and Maicer Izturis both injured.
"This guy can play the game," Scioscia said.
Rodriguez is batting .290 at Salt Lake with 23 homers and 79 RBIs in 81 games. Wood is batting .316 in the PCL with 17 homers and 55 RBIs in 74 games, while Terry Evans checks in at .290 with 22 homers, 76 RBIs in 105 games. Rodriguez has a narrow edge over Wood in slugging, .603 to .597, with Evans at .526.
Recalling fatherly wisdom
When Gary "Sarge" Matthews was teaching his son the finer points of the game during Gary Jr.'s youth, there were insights and expressions culled from a life spent in baseball that resonate all these years later with the Angels' outfielder.
"My dad used to tell me that he could teach me how to hit breaking balls and changeups – but either you can hit a fastball or you can't," Gary Jr. was saying on Father's Day, his dad in Philadelphia where he works as a commentator on Phillies telecasts.
"My dad is as old school as it gets. That still rings true, but I have formed my own opinions over the years. I think one change in the game is that pitchers have evolved and now throw more breaking balls for strikes. Controlling the breaking ball, and not relying on fastballs as much early in counts, has changed things."
In Detroit on the recent road trip, Matthews launched a 100-mph heater by Joel Zumaya – the hardest thrower in the game – deep into the right-field seats, foul. He'd turned on triple digits and was a split-second out in front of it, a display of remarkably quick hands.
On Saturday night at Angel Stadium, Matthews came off the bench in the ninth inning and launched a fastball from Dodgers reliever Jonathan Broxton into the seats in right center, turning a 6-2 deficit into what would be a 6-4 loss. It was the first Angels pinch-hit homer of the year and the fourth of Matthews' career.
"I've never been accused of not being able to hit a fastball," Matthews said, grinning.
Walking past Matthews' locker, hitting coach Mickey Hatcher said, "You can't put one past him."
Matthews is a man without a position, a man who wants to play every day but has no steady job with the Angels. Juan Rivera has been on fire in left, joining Torii Hunter and Bobby Abreu in the outfield with Vladimir Guerrero absorbing designated-hitter at-bats.
Matthews has made it clear all season that he doesn't think of himself as a backup and will not be happy in that role. He thinks he's one of the game's most gifted center fielders, and Hunter – the best – agrees. But that is not much consolation. When you're an athlete and you're sitting, you don't feel right.
All Matthews can do now is accept his role and make the best of a difficult situation. He is an expensive insurance policy, a card that will remain in manager Mike Scioscia's deck most of the time until someone in front of him is injured or falls into a major slump.
"Not much I can do about it," Matthews said. "I'll just keep working and be ready when I'm called on."
His old-school dad is on the cellphone with his son all the time, offering perspective, support, all the things a young man needs when he's frustrated.
It is possible something could happen around the July 31 Non-Waiver Trade Deadline, Matthews acquiring sudden appeal to a club in need of a quality center fielder. His salary – he's in the third year of a five-year, $50 million contract – makes it unlikely. And not because Matthews has the contractual right to refuse a trade, as Jake Peavy did when the White Sox and Padres had worked out a deal.
Matthews yearns to be an everyday center fielder, but he's on a club that employs the game's best. It's like being the guy who thought he'd have a crack at the ceiling of the Sistine Chapel before Michelangelo showed up with his brushes and paints.


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So while the fire department puts out the gas boiler thats on fire over the road a thought crossed my mind. Why dont we have more LCARS keyboards? The traditional keyboard layout is adopted from type writers because when they where common place, it was the only layout that stopped the character hammers from getting tangled when typing relatively fast. The layout was translated into the PC world for familiarity and ease of transition. But lets face it, the world changes. Voice recognition software is coming along and keyboards are becoming less and less important. So why not compact them, and change them to something more versatile like the LCARS type controls? Sure, you can still use your generic old school keyboard but its time for a change and the entire way computers are being used is leaning more and more towards touch interfaces, so the traditional keyboard is soon going to find itself in a world where its no longer required. Why not make that change sooner?
thats the voice recognitions that take over, you wouldn't need to type really, just dictate. Or the pads could just change mode from command type to text type, just a different layout.
It wasn't just for that. The "QWERTY" format was also adopted because of its practicality in the office world. For skilled typists, the placement of letters allows words to be typed with the left hand, while the right hand has access both to the cursor buttons and the numeric keypad. Primarily because LCARS is just made up stuff for a TV show. Our heroes can do all sorts of stuff on LCARS interfaces, including typing text and codes with buttons marked only with tiny little numbers due to magic of TV. But as far as keyboards becoming less and less important, I would argue that they would still be important as long as there's a need to write messages or text to people. They may disappear when the need for people to write does, though.
There was a discussion on this topic some months ago. If you're going to type text with any degree of speed or efficiency, you need the tactile sensation of pressing individual keys. Touch screens just won't do the job. The traditional QWERTY keyboard layout has been around for more than 130 years, and it's not going to become obsolete anytime soon.
Voice recognition software currently is still pretty awful. Actually, I wonder why the helm isn't outfitted with a mouse.
In work in a newsroom with fifteen other people. Do you really think we should dictate everything we write out loud?
But I can type faster than I can talk. Plus voice recognition software can't seem to handle my accent.
And if the computer can properly recognize your voice, what possible purpose could a keyboard of any sort serve? If you don't need a button for the letter G, why would you need a button for anything else, either? As for touchscreens vs. tactile feedback, there's no reason the two should be mutually exclusive. Indeed, a keyboard where the keys are not physically protruding objects as such, but mere virtual creations with pseudo-physical characteristics (such as buzzing your finger when you press), would be advantageous in having the ability to move the key for letter G beneath your finger even when you are in danger of missing it. Timo Saloniemi
Interestingly, they were far ahead of their time in the TOS pilots. In "The Cage" / "The Menagerie" Spock was doing his "power point" presentation of the Talos star group by the move of his right hand without touching a panel or a button. And in "Where No Man Has Gone Before" Gary Mitchell just activated a component on his console by letting his hand pass over the console without ever touching it. Unfortunately, in "The Corbomite Maneuver" they went back to the traditional pushing of buttons. Bob
You misunderstand - I wanted to postulate a hypothetical situation where voice recognition (or, say, telepathic recognition) works with absolute perfection, come hell or high water or the noises of a newsroom. It now perfectly takes your dictation. What then? The OP seems to want an input device that does away with the inputting of text by typing. Well, now he has got it. What possible use could he now have for a keyboard, be it a dull QWERTY one or a flashy LCARS one? Everything he could ever want input can now be input the same way his keyboard-sidestepping dictation can. Timo Saloniemi
Then we have fifteen people talking at the same time in the same tiny room, while trying to concentrate just enough to write complex articles, which by the way is not easy to do when you have to dictate them. It's not a matter of technology. It's a matter of being able to think.
And still the point stands - no matter whether the clearing of those hurdles is possible or not, it's simply not going to produce the result the OP wanted. Designing a machine capable of taking your dictation is possible one way or another, make no mistake, but yeah, it may be somewhat difficult. Yet when it finally happens, the one thing it won't result in is a fancy new type of keyboard. Timo Saloniemi
I can see why they went back to traditional button-pushing when the show went into production. The hand-waving made Spock and Gary Mitchell look like stage magicians.
I have to add that the voice recognition software of the Google app on my Surface is freakishly efficient.
To be sure, neither of the sleighs-of-hand may necessarily have been intended to be interpreted that way. It's well-known that Spock in that scene is gesturing next to an observant seated underling who probably was intended to be pushing the buttons of the slide projector, but very fortunate framing of the shot leaves the gal out of the picture and creates this far more futuristic impression. Gary in turn simply seems to prefer punching the buttons with the edge of his hand, rather than his fingers, as this appears more laid-back in keeping with his character. Timo Saloniemi

subset: commoncrawl


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A remarkable diversity of bone-eating worms (Osedax; Siboglinidae; Annelida)
- Robert C Vrijenhoek1Email author,
- Shannon B Johnson1 and
- Greg W Rouse2
© Vrijenhoek et al; licensee BioMed Central Ltd. 2009
Received: 29 June 2009
Accepted: 10 November 2009
Published: 10 November 2009
Bone-eating Osedax worms have proved to be surprisingly diverse and widespread. Including the initial description of this genus in 2004, five species that live at depths between 25 and 3,000 m in the eastern and western Pacific and in the north Atlantic have been named to date. Here, we provide molecular and morphological evidence for 12 additional evolutionary lineages from Monterey Bay, California. To assess their phylogenetic relationships and possible status as new undescribed species, we examined DNA sequences from two mitochondrial (COI and 16S rRNA) and three nuclear genes (H3, 18S and 28S rRNA).
Phylogenetic analyses identified 17 distinct evolutionary lineages. Levels of sequence divergence among the undescribed lineages were similar to those found among the named species. The 17 lineages clustered into five well-supported clades that also differed for a number of key morphological traits. Attempts to determine the evolutionary age of Osedax depended on prior assumptions about nucleotide substitution rates. According to one scenario involving a molecular clock calibrated for shallow marine invertebrates, Osedax split from its siboglinid relatives about 45 million years ago when archeocete cetaceans first appeared and then diversified during the late Oligocene and early Miocene when toothed and baleen whales appeared. Alternatively, the use of a slower clock calibrated for deep-sea annelids suggested that Osedax split from its siboglinid relatives during the Cretaceous and began to diversify during the Early Paleocene, at least 20 million years before the origin of large marine mammals.
To help resolve uncertainties about the evolutionary age of Osedax, we suggest that the fossilized bones from Cretaceous marine reptiles and late Oligocene cetaceans be examined for possible trace fossils left by Osedax roots. Regardless of the outcome, the present molecular evidence for strong phylogenetic concordance across five separate genes suggests that the undescribed Osedax lineages comprise evolutionarily significant units that have been separate from one another for many millions of years. These data coupled with ongoing morphological analyses provide a solid foundation for their future descriptions as new species.
Osedax, a recently discovered genus of bone-eating marine worms, are proving to be far more diverse and geographically widespread than initially realized. The genus was described from two newly discovered species found on whalebones recovered from 2,893 m depth in Monterey Bay, California [1]. Subsequently, three additional species were described from depths between 30 and 3,000 m in the Atlantic and Pacific oceans [2–4]. Now, five additional distinct evolutionary lineages are recognized from Monterey Bay, but these putative species remain to be formally described [5–8]. Here we report genetic evidence for seven additional putative species. Given this unexpected diversity of Osedax worms with distinct morphologies, depth ranges and ecological characteristics, a detailed examination of their evolutionary history is warranted.
The initial description of Osedax [1] included a phylogenetic analysis that placed the new genus in the polychaete annelid family Siboglinidae, which also includes the now obsolete tubeworm phyla Vestimentifera and Pogonophora [9, 10]. As adults, all siboglinids lack a functional digestive system and rely entirely on endosymbiotic bacteria for their nutrition. The other siboglinid taxa host chemosynthetic bacteria and live in reducing marine environments such as hydrothermal vents, hydrocarbon seeps and anoxic basins. Osedax, however, are unique because they penetrate and digest bones with the aid of heterotrophic bacteriathat are housed in a complex branching root system [6, 11]. Osedax also differ because they exhibit extreme sexual dimorphism involving dwarf (paedomorphic) males that live as harems within a female's tube [1, 4, 12].
Characteristics of Osedax OTUs.
Taxa | Authority1 | Depth (m) | Approx. Size2 | Palp color | Pinnules3 | Oviduct4 | Roots | Museum Voucher5 |
O. rubiplumus | Rouse et al. 2004 | 1820--2893 | 59 | brilliant red | out | long | long, branched | CASIZ 170238 |
O. frankpressi | Rouse et al. 2004 | 1820--2893 | 23 | red with white lateral stripe | in | long | robust, lobate | CASIZ 170235 |
O. mucofloris | Glover et al. 2005 | 30--125 | 14 | white to pink | in | long | robust, lobate | |
O. japonicus | Fujikura et al. 2006 | 224--250 | 16 | pink | out | short | robust, lobate | |
O. roseus | Rouse et al. 2008 | 633--1820 | 24 | bright red | out | long | long, branched | SIO-BIC A979-981 |
O. spiral | Braby et al. 2007 | 2893 | 25 | no palps | none | absent | long thin filaments | SIO-BIC A1639 |
O. yellow-collar | Braby et al. 2007 | 383 | 18 | red | both | long | robust, lobate | SIO-BIC A1640 |
O. orange-collar | Braby et al. 2007 | 383--1018 | 18 | red | both | long | robust, lobate | SIO-BIC A1641 |
O. nude-palp-A | Jones et al. 2008 | 1820 | 25 | red | none | ? | ? | SIO-BIC A1642 |
O. nude-palp-B | Jones et al. 2008 | 2893 | 25 | red | none | ? | ? | SIO-BIC A1643 |
O. nude-palp-C | Rouse et al. 2009 | 1018 | 12 | red | none | ? | ? | SIO-BIC A1644 |
O. nude-palp-D | this report | 1018-1820 | 12 | red | none | ? | ? | SIO-BIC A1645 |
O. nude-palp-E | this report | 1018 | 12 | red | none | ? | robust, lobate | SIO-BIC A1646 |
O. nude-palp-F | this report | 2892 | 18 | red | none | ? | robust, lobate | SIO-BIC A1647 |
O. white-collar | this report | 1018 | 6 | red with white stripe | both | long | robust, lobate | SIO-BIC A1648 |
O. yellow-patch | this report | 633--1018 | 5 | pale | both | long | robust, lobate | SIO-BIC A1649 |
O. green-palp | this report | 1820 | 3 | red/green | ? | long | robust, lobate | SIO-BIC A1650 |
S. brattstromi | Webb 1964 | 600 | 40 | red | none | absent | none |
Characterization of DNA sequences and the substitution models used to correct for saturation in the Bayesian analyses.
DNA | Length | Mean % divergence1 | % GC content | Ts:Tv2 | Model | ||
Protein-coding | Synonomous3 | Nonsynonymous | |||||
mt COI | 1005 | 17.0 | 41.0 | 1.304 | 259.42 | 94.58 | GTR+SS |
nuc H3 | 371 | 13.9 | 48.8 | 1.075 | 222.23 | 71.77 | Sym+SS |
Ribosomal | Indels4 | Size5 | |||||
mt 16S | 506 | 11.1 | 32.6 | 0.527 | 3 | 1.0 | GTR+I+G |
nuc 18S | 1644 | 5.9 | 48.0 | 0.526 | 5 | 1.18 | GTR |
nuc 28S | 400 | 8.6 | 47.4 | 0.584 | 4 | 1.14 | GTR+I+G |
Phylogenetic analyses
COI sequence divergence (K2P corrected) within (π in italics on diagonal) and among (D on lower left) the Osedax OTUs.
1 | O. rubiplumus | 0.0051 | ||||||||||||||||
2 | O. frankpressi | 0.1934 | 0.0034 | |||||||||||||||
3 | O. roseus | 0.1573 | 0.1836 | 0.0047 | ||||||||||||||
4 | O. white-collar | 0.1957 | 0.1985 | 0.2048 | 0.0017 | |||||||||||||
5 | O. pinnules | 0.2136 | 0.2131 | 0.2159 | 0.2350 | 0.0070 | ||||||||||||
6 | O. green-palp | 0.1795 | 0.1869 | 0.1859 | 0.2124 | 0.1952 | 0.0020 | |||||||||||
7 | O. yellow-collar | 0.1896 | 0.1963 | 0.1831 | 0.1870 | 0.1897 | 0.2073 | 0.0048 | ||||||||||
8 | O. orange-collar | 0.1838 | 0.2018 | 0.1844 | 0.1821 | 0.2018 | 0.2042 | 0.0843 | 0.0078 | |||||||||
9 | O. spiral | 0.2028 | 0.2069 | 0.2017 | 0.2148 | 0.2016 | 0.2086 | 0.1999 | 0.1990 | 0.0019 | ||||||||
10 | O. nude-palp-A | 0.2025 | 0.1929 | 0.2034 | 0.2354 | 0.2091 | 0.1879 | 0.2013 | 0.2016 | 0.2001 | 0.0082 | |||||||
11 | O. nude-palp-B | 0.2028 | 0.1919 | 0.1902 | 0.2216 | 0.2181 | 0.1996 | 0.2166 | 0.2298 | 0.1817 | 0.1674 | 0.0000 | ||||||
12 | O. nude-palp-C | 0.1899 | 0.1927 | 0.2001 | 0.2312 | 0.2057 | 0.2039 | 0.2188 | 0.1964 | 0.1966 | 0.1707 | 0.1593 | 0.0060 | |||||
13 | O. nude-palp-D | 0.1950 | 0.2204 | 0.1971 | 0.2367 | 0.2140 | 0.2054 | 0.2339 | 0.2191 | 0.2063 | 0.1956 | 0.1888 | 0.1643 | 0.0141 | ||||
14 | O. nude-palp-E | 0.2110 | 0.2250 | 0.2279 | 0.2087 | 0.2029 | 0.2228 | 0.2073 | 0.2176 | 0.1984 | 0.1926 | 0.1982 | 0.1713 | 0.1880 | 0.0072 | |||
15 | O. nude-palp-F | 0.1781 | 0.2100 | 0.1909 | 0.1933 | 0.2072 | 0.1948 | 0.2084 | 0.2114 | 0.1919 | 0.1823 | 0.1643 | 0.1626 | 0.1822 | 0.1723 | - | ||
16 | O. mucofloris | 0.2341 | 0.2106 | 0.2063 | 0.2138 | 0.2241 | 0.2432 | 0.1855 | 0.1856 | 0.2001 | 0.2075 | 0.2305 | 0.2061 | 0.2362 | 0.1950 | 0.2355 | 0.0060 | |
17 | O. japonicus | 0.2064 | 0.2125 | 0.1934 | 0.1788 | 0.2082 | 0.2145 | 0.1602 | 0.1426 | 0.2156 | 0.2215 | 0.2095 | 0.2307 | 0.2431 | 0.2378 | 0.2203 | 0.1726 | - |
Four additional genes revealed concordant phylogenetic differences among the Osedax OTUs (Figures 2b-e). The 16S, 28S, and H3 sequences differed among the 15 Monterey Bay OTUs, but sequences were not available for O. japonicus and O. mucofloris. Although their 18S sequences were identical, O. yellow-collar and O. orange-collar differed from all the other Monterey OTUs and from O. mucofloris. The nodes leading to O. spiral and O. frankpressi were not stable, but all five gene-trees were broadly congruent in their topologies. Incongruence length difference (ILD) tests of homogeneity revealed that four of the five gene partitions were not significantly in conflict (P range: 0.119 - 1.00). Only the H3 tree was incongruent with respect to the 16S and 18S rRNA trees (P = 0.03 and 0.02, respectively). This problem resulted because H3 provided weak resolution among Osedax species that clustered together on a long branch relative to the outgroup S. brattstromi. ILD tests of homogeneity between H3 and the other partitions without the outgroup eliminated all remaining incongruence (P range: 0.125 - 1.00).
Species diversity
Genetic and morphological differences among five previously named Osedax species provide a useful reference frame for assessing levels of divergence among the twelve undescribed OTUs considered in this study. Osedax rubiplumus, O. frankpressi and O. roseus live together on whale carcasses at depths greater than 1,000 m in Monterey Bay, CA (Figure 1; Table 1). To date, we have found no evidence for interbreeding among them. For example, an examination of 116 male Osedax sampled from the tubes of 77 O. rubiplumus females found no cases of foreign males in the female's tubes, despite the presence of O. roseus and O. frankpressi on the same carcass at 1820 m depth. Also including O. mucofloris from Sweden and O. japonicus from Japan, the mean sequence divergence (D) for mitochondrial COI between pairs of the named species was 19.6% (range: 15.7 to 23.4%; Table 2). Correspondingly, the mean pairwise D among the undescribed OTUs was 19.9% (range: 8.4 to 23.7%). The smallest value (8.4% between OTUs O. yellow-collar and O. orange-collar) was an order-of-magnitude greater than the largest divergence observed within any of these named or undescribed OTUs (π = 0.8% for O. nude-palp-A). These π values probably are underestimates, however, because each was obtained from a single locality. Isolation-by-distance and population subdivision across oceanic barriers are expected to increase π within broadly distributed species; however, π rarely exceeds 1 - 2%, unless other factors are involved. Global-scale phylogeographic surveys of COI sequence diversity have estimated π values less than 1% within named species of deep-sea hydrothermal vent annelids, mollusks and crustaceans, whereas D values typically are greater than 4% among species [14–22]. Nonetheless, odd cases of accelerated COI substitution rates have been reported. Sex-biased mitochondrial transmission and heteroplasmy are associated with accelerated divergence in some bivalve mollusks [23], but no evidence exists for this phenomenon in annelids, and we have found no differences in the distribution of mitochondrial haplotypes between males and females for O. rubiplumus [12]. High mitochondrial divergence rates have been reported for some marine and freshwater animals [24–26], but for the vast majority of cases mitochondrial COI is relatively conservative in its mutation rate within and among species. It is precisely the tendency of COI to discriminate clearly among the named species in many invertebrate taxa that has made this gene a common reference tool for DNA barcoding and molecular taxonomy [27, 28].
GenBank accession numbers for the DNA sequences used in this study.
Taxa | COI | 16S | 18S | 28S | H3 |
O. rubiplumus | EU223307--08, EU223298--99, DQ996618, 20 | FJ347656 | FJ347681 | FJ347671 | FJ347704 |
O. frankpressi | AY586495, EU223314, DQ996621, FJ347605--07 | FJ347658 | FJ347682 | FJ347674 | FJ347705 |
O. mucofloris | AY827562-568 | AY941263 | |||
O. japonicus | AB259569 | ||||
O. roseus | EU164760--61, EU032469--70, FJ347607-08 | FJ347657 | FJ347683 | FJ347670 | FJ347709 |
O. spiral | DQ996622--24, FJ347636--38 | FJ347647 | FJ347693 | FJ347676 | FJ347703 |
O. yellow-collar | DQ996629, 32--33, EU223332--33, EU223335 | FJ347660 | FJ347689 | FJ347672 | FJ347706 |
O. orange-collar | EU223340--41, EU223354, FJ347627-29 | FJ347661 | FJ347690 | FJ347673 | FJ347707 |
O. nude-palp-A | EU223356--58, FJ347622--24 | FJ347653 | FJ347687 | FJ347662 | FJ347702 |
O. nude-palp-B | EU236218 | FJ347652 | FJ347686 | FJ347665 | FJ347701 |
O. nude-palp-C | EU267675--76, J347625--26 | FJ347650 | FJ347688 | FJ347666 | FJ347710 |
O. nude-palp-D | FJ347630--31 | FJ347649 | FJ347691 | FJ347667 | FJ347708 |
O. nude-palp-E | FJ347632--35 | FJ347648 | FJ347692 | FJ347664 | FJ347700 |
O. nude-palp-F | FJ347643 | FJ347651 | FJ347695 | FJ347663 | FJ347699 |
O. white-collar | FJ347610--15 | FJ347659 | FJ347684 | FJ347675 | FJ347712 |
O. yellow-patch | FJ347616--21 | FJ347654 | FJ347685 | FJ347668 | FJ347698 |
O. green-palp | FJ347639--42 | FJ347655 | FJ347694 | FJ347669 | FJ347711 |
S. brattstromi | FJ347645 | FJ347680 | FJ347677 | FJ347697 | FJ347645 |
Individual gene trees (Figure 2) and the combined phylogenetic analysis (Figure 3) identified several well-supported groupings within Osedax (clades I - V). Osedax spiral (clade III) stands alone as the most atypical of these worms. Its oviduct does not extend beyond the trunk, and it lacks the vascularized anterior palps that characterize all other Osedax (Figure 1g). Unlike all other Osedax, O. spiral is a late successional species which lives at the sediment interface and produces long fibrous roots that penetrate the anoxic (black and sulfidic) sediments to exploit buried fragments of bone [5]. The lack of palps in O. spiral probably represents a character loss under a most parsimonious reconstruction, because all other siboglinids bear an anterior crown composed of one or more palps. The nude-palp OTUs (clade II) differ because their palps do not bear the lateral pinnules seen in the other Osedax clades (Figures 1i and 1j). Lack of pinnules may represent a character loss, but supporting evidence regarding the homology and distribution of pinnules in other siboglinids is uncertain. Monoliferans have two or more palps, with numerous pinnules in the case of Vestimentifera, but pinnules are absent in Sclerolinum and in some frenulates [10, 35].
The remaining Osedax clades (I, IV and V) bear four palps with numerous pinnules that give the crown a feathery appearance (e.g., Figure 1a). The two members of clade V have long branched roots that are green in color (Figures 1e-f) and palps that are bright red with outwardly facing pinnules. Clades II and V share robust lobate roots. The two members of clade (I) have relatively short trunks and palps (Figure 1h), but they have not been found in great numbers because they are small and may have been overlooked in earlier samples. Members of clade IV, which have red, pink or even white crowns (Figures 1a-d), were recovered from depths of 1,020 m or less, excepting O. frankpressi, which has not been found shallower than 1,800 m. Occupation of shallow habitats might be a derived condition for these members of clade IV, though support for a shallow clade was weak (Figure 3). Addition of comparative sequence data from the other shallowmembers of this clade, O. japonicus and O. mucofloris, might help to strengthen this relationship (only 18S and COI data are available on GenBank for O. mucofloris and only COI for O. japonicus). Otherwise, no clear evolutionary pattern of depth utilization is apparent among the major Osedax clades. Several of these OTUs were sampled from a single depth, others were sampled across relatively narrow depth ranges (300 - 600 m for O. yellow-patch and O. orange-collar), and some were sampled across broad depth ranges (1,000 m for O. frankpressi, and O. rubiplumus and 1,200 m for O. roseus).
Age of Osedax
For now, we are unable to confidently delineate a timeframe during which Osedax split from its monoliferan relatives or the age (T) of the most recent common ancestor for this unusual genus. Present evidence indicates that Osedax species live primarily on organic compounds extracted directly from sunken bones. Their Oceanospirillales symbionts are capable of growing on collagen and cholesterol as primary carbon sources [6]. Video evidence suggests that O. japonicus also grows on spermaceti, a wax found in the head of sperm whales [3]. There are arguments to suggest that Osedax may not be nutritionally restricted to living on whale-falls. Experimental deployments of cow bones and observations of sunken possible pig bones reveal that Osedax can grow and reproduce on a range of mammalian tissues including those from terrestrial quadrupeds [7, 36]. So, it may be unwarranted to associate the evolution of these bone-eating worms with the origin and spread of oceanic whales, as previously suggested by Rouse et al. [1]. Nonetheless, one of the scenarios that we have considered here is consistent with that hypothesis. If we assumed a divergence rate (d = 1.4% per MY) calibrated for mitochondrial genes from shallow-water marine invertebrates [13], and applied this rate (r 1 = d/2 = 0.70%/lineage/My) to COI divergence, we estimated that Osedax split from its monoliferan relatives about 45 Mya, possibly coincident with the origins of large archeocete cetaceans during the Eocene [37]. According to this scenario, the most recent common ancestor for the Osedax sampled to date would have lived about 26 MYA, during the Late Oligocene and roughly coincident with the diversification of modern cetaceans [38].
Alternatively, we can assume a slower substitution rate (r 1 = 0.21%/lineage/My) calibrated from COI divergence in deep-sea annelids, including Vestimentifera [14], as was used by Rouse et al. [1] for estimating the origin of Osedax when only O. rubiplumus and O. frankpressi were known. Under this rate then Osedax would appear to be much older than previously hypothesized [1]. This result is not surprising given the larger diversity of Osedax shown here. Accordingly, Osedax split from its monoliferan relatives during the Cretaceous, and the most recent common ancestor for the genus would have lived during the Late Cretaceous. Perhaps the calcified cartilage and bones from a variety of large Cretaceous vertebrates supported these worms -- e.g., mosasaurs, plesiosaurs, turtles, and possibly chondrichthyans and teleosts [39–42]. Fossilized snails and bivalves were recently found with plesiosaur bones; so the sunken carcasses of these large marine reptiles appear to be capable of supporting communities much like those found on modern whale-falls [43]. Nevertheless, this scenario is problematic, because the major Osedax clades would have diversified around the Cretaceous-Tertiary (K/T) boundary, after the extinction of most large-bodied reptilians [44]. Although dyrosaurid crocodylomorphs survived the K/T event, they were confined to relatively shallow coastal environments [45] and probably would not have supported Osedax. Large turtles and chondrichthyans also survived the K/T boundary [42], and large teleosts appeared again during the early Paleocene [46]. It is unknown whether Osedax can exploit these resources; so, arguably a 20 MY gap may have existed during the Paleocene when there would have been little in the way of large vertebrate remains for Osedax. Another problem with this scenario is the concern that nucleotide substitution rates may be slower in the deep-sea vent annelids used to obtain the r 2 = 0.21% calibration rate [9].
The present phylogenetic evidence based on DNA sequences from multiple independent genes provides a solid foundation for future discoveries and taxonomic descriptions of Osedax species. However, our efforts to estimate evolutionary ages for the diversification of this unusual group of worms only allowed the erection of new hypotheses that could be tested with independent evidence from the fossil record. Soft-bodied invertebrates like Osedax do not often leave convincing fossils, but these worms might leave traces of their activity by the distinctive holes they bore into bones. To date, we have found no other animals that create similar borings in bones. Consequently, we have distributed whalebones containing Osedax to several paleontologists who are also examining the taphonomy of fossilized bones from plesiosaurs and cetaceans. It is to be hoped that these efforts will help us to narrow the age of this remarkable genus of bone-eating worms.
Materials and methods
Locations of the Monterey Bay whale-falls, except whale-634, are provided elsewhere [5]. Whale-634 is the carcass of a juvenile gray whale that was sunk on 5 October 2004 at a depth of 633 m at 36.802°N and 122.994°W. We used the remotely operated vehicles, ROV Tiburon and ROV Ventana, operated by the Monterey Bay Aquarium Research Institute (MBARI) to collect Osedax-inhabited bones from five whale-fall localities (Table 1). Bones were transported to the surface in closed insulated containers and stored temporarily in cold (4°C) filtered seawater. Worms were dissected from the bones and photographed. Then a palp tip was removed and stored in 95% ethanol or frozen immediately at -80°C. The remainder of each specimen was preserved for anatomical studies and taxonomic descriptions. Voucher specimens were lodged in Scripps Institution of Oceanography Benthic Invertebrate Collection (catalogue numbers in Table 1). Other specimens will be distributed to other Museums upon their formal description (Rouse, in progress). For the present purpose, we list the approximate sizes (trunk plus crown length) and several morphological characteristics of each OTU (Table 1).
Published DNA sequences from Osedax mucofloris (18S rRNA and COI) and O. japonicus (COI) were recovered from GenBank [2]. A previous phylogenetic analyses [1] placed Osedax in a clade that also includes the Monolifera, which includes Sclerolinum and vestimentiferan tubeworms [10]. The frenulates, a diverse group of slender chemosynthetic worms are basal to the monoliferans and Osedax [10, 47]. Ongoing studies of siboglinid phylogeny revealed that Sclerolinum is presently our best choice as outgroup for this study of Osedax phylogeny. The vestimentiferan Lamellibrachia columna was also examined, and its substitution as outgroup did not substantively alter the tree topologies for the ingroup. Other vestimentiferans were not considered, however, because incomplete sequence data are available and because independent evidence from several genes suggests that rates of nucleotide substitution may have slowed down in these deep-sea worms [9, 14, 48]. Consequently, we have used DNA sequences from the monoliferan Sclerolinum brattstromi, collected near Bergen, Norway. GenBank accession numbers for all the DNA sequences used in this study are listed in Table 4.
DNA methods
Total DNA was extracted using the DNeasy kit (Qiagen, Valencia, CA, USA) according to manufacturer's instructions. We used primers that amplified approximately 1200 bp of COI [49], approximately500 bp of 16S rRNA [50], approximately 1000 bp of 28S rRNA [51], approximately 1800 bp of 18S rRNA [52], and approximately 370 bp of H3 [53]. Amplification reactions with AmpliTaq Gold (Applied Biosystems Inc., Foster City, CA, USA) were conducted in a GeneAmp 9700 thermal cycler (Applied Biosystems Inc., Carlsbad, CA, USA) with the following parameters: 95°C/10 min, 35× (94°C/1 min, 55°C/1 min, 72°C/1 min), and 72°C/7 min. If available, at least six individuals of each species were sequenced for each locus. PCR products were diluted in 50 μl sterile water and cleaned with Multiscreen HTS PCR 96 filter plates (Millipore Corp., Billerica, MA, USA). The products were sequenced bidirectionally with the same primers on an ABI 3100 sequencer using BigDye terminator v.3.1 chemistry (Applied Biosystems Inc., Foster City, CA, USA).
Phylogenetic analyses
Sequences were assembled using CodonCode Aligner v. 2.06 (CodonCode Corporation, Dedham, MA, USA), aligned using Muscle [54] and edited by eye using Maclade v. 4.08 [55]. We used MrModelTest [56] and the Akaike information criterion [57] to determine appropriate evolutionary models for each gene (Table 2). COI and H3 were partitioned by codon position, and parameters were estimated separately for each position. RNA secondary structures were predicted with GeneBee and used to partition stems and loops in 16S, 18S, and 28S sequences. The doublet model was used for RNA stems and a standard 4 × 4 nucleotide model was used for RNA loops. The number of indel haplotypes for rRNA sequences (total number of indels, number after excluding overlapping indels, and average length of indels) were estimated with DNAsp v. 4.90.1 [58] using the diallelic model. Gaps in the RNA sequences were treated as a fifth character-state in subsequent Bayesian phylogenetic analyses and as missing data in parsimony and maximum likelihood (ML) analyses. The program DAMBE [59] was used to examine saturation of the mitochondrial COI sequences for the Osedax OTUs and outgroup taxa.
First, each gene was analyzed separately using MrBayes v. 3.1.2 [60, 61]. Bayesian analyses were run as six chains for 5·106 generations. Print and sample frequencies were 1,000 generations, and the burn-in was the first 100 samples. We used AWTY [62] to assess whether analyses reached convergence and FigTree v. 1.1.2 [63] to display the resulting trees. We then used the incongruence length difference (ILD) function implemented in Paup* v. 4.0 [64] to assess congruence of the tree topologies produced by the individual gene partitions. ILD tests were conducted both with and without the outgroup taxa. The ILD partition homogeneity test was run for 1,000 replicates with 10 random additions of gene sequences.
A combined analysis was conducted with concatenated sequences from the five genes. If available, multiple individuals of each OTU were sequenced for each gene; however, the concatenated multilocus sequences used in the phylogenetic analyses were obtained from a single representative individual for each OTU. The five gene regions were partitioned separately according to the previously determined model parameters. Bayesian phylogenetic analyses were then conducted with MrBayes v. 3.1.2. Maximum parsimony analysis of the combined data set was performed with Paup* v. 4.0 [64] using an equally weighted character matrix, heuristic searches using the tree-bisection-reconnection branch-swapping algorithm, and 100 random addition replicates. The resulting shortest tree included 3481 steps. A parsimony jackknife analysis (with 37% deletion) was run for 100 iterations with the same settings as the parsimony search. ML analysis was conducted using RAxML 7.0.4 (with bootstrapping) using GTR+I+G as the model for each partition on combined data. RAxML analyses were performed with the CIPRES cluster at the San Diego Supercomputer Center.
Relaxed molecular clock
A Bayesian, MCMC method implemented in Beast v. 1.4.8 [65] was used to estimate the evolutionary ages of internal nodes in the tree topology derived from the combined phylogenetic analysis. Estimates of the time to most recent common ancestor (T) were based on two calibrations nucleotide substitution rates for mitochondrial COI. Substitution rates (r) were estimated as percentage per lineage per million years (my) so they equal one-half the divergence per unit of time (T) between taxa (r = 100 × D/2T). First, we assumed a conventional substitution rate, r 1 = 0.7%, based on D = 1.4% per my pairwise divergence rate commonly cited for shallow water marine invertebrates that were isolated by the emergence of the Isthmus of Panama [13]. Second, we used a slower rate, r 2 = 0.21%, previously calibrated from a vicariant event that split cognate-species of deep-sea hydrothermal vent annelids between the East Pacific Rise and the northeastern Pacific ridge system about 28.5 myA [14]. Calibrations were not available for the other genes.
We used a relaxed, uncorrelated, lognormal molecular clock with a general time reversible (GTR) substitution model that was unlinked across codon positions. Initial MCMC test runs consisted of 10 million generations to optimize the scale factors of the prior function. Three independent MCMC chains were run for 100 million generations, sampled every 1000 generations. Results were visualized in and FigTree v. 1.1.2 and Tracer v. 1.4 [66].
mitochondrial large subunit ribosomal RNA
nuclear small subunit subunit ribosomal RNA
nuclear large subunit ribosomal RNA
cytochrome oxidase subunit I
general time reversible
Histone 3
Monte Carlo Markov chain
million (106) years ago
- OTUs:
operational taxonomic units
remotely operated vehicle.
We thank the captains and crews of the R/V Western Flyer and R/V Point Lobos and the pilots of ROV Tiburon and ROV Ventana for their technical support and patience. We gratefully appreciate the role that W. Joe Jones played in the early phases of this research. Thanks to Ken Halanych for providing a tissue sample from Sclerolinum brattstromi, and to Julio Harvey for his editorial help. Funding for this project was provided by the Monterey Bay Aquarium Research Institute (The David and Lucile Packard Foundation), NSF Award #0334932 (under the Assembling the Tree of Life program) to investigate the Tree of Life for Protostomes, and SIO for start up funds to GWR.
Authors' Affiliations
- Rouse GW, Goffredi SK, Vrijenhoek RC: Osedax: Bone-eating marine worms with dwarf males. Science. 2004, 305: 668-671. 10.1126/science.1098650.View ArticlePubMedGoogle Scholar
- Glover AG, Käilström B, Smith CR, Dahlgren TG: World-wide whale worms? A new species of Osedax from the shallow north Atlantic. Proc R Soc Lond B. 2005, 272: 2587-2592. 10.1098/rspb.2005.3275.View ArticleGoogle Scholar
- Fujikura K, Fujiwara Y, Kawato M: A new species of Osedax (Annelida: Siboglinidae) associated with whale carcasses off Kyushu, Japan. Zoolog Sci. 2006, 23: 733-740. 10.2108/zsj.23.733.View ArticlePubMedGoogle Scholar
- Rouse GW, Worsaae K, Johnson SB, Jones WJ, Vrijenhoek RC: Acquisition of dwarf male 'harems' by recently settled females of Osedax roseus n. sp. (Siboglinidae; Annelida). Biol Bull. 2008, 214: 67-82. 10.2307/25066661.View ArticlePubMedGoogle Scholar
- Braby CE, Rouse GW, Johnson SB, Jones WJ, Vrijenhoek RC: Bathymetric and temporal variation among Osedax boneworms and associated megafauna on whale-falls in Monterey Bay, California. Deep Sea Res I. 2007, 54: 1773-1791. 10.1016/j.dsr.2007.05.014.View ArticleGoogle Scholar
- Goffredi SK, Johnson SB, Vrijenhoek RC: Genetic and potential function of microbial symbionts associated with newly discovered species of Osedax polychaete worms. Appl Environ Microbiol. 2007, 73: 2314-2323. 10.1128/AEM.01986-06.PubMed CentralView ArticlePubMedGoogle Scholar
- Jones WJ, Johnson SB, Rouse GW, Vrijenhoek RC: Marine worms (genus Osedax) colonize cow bones. Proc R Soc Lond B. 2008, 275: 387-391. 10.1098/rspb.2007.1437.View ArticleGoogle Scholar
- Rouse GW, Wilson NG, Goffredi SK, Johnson SB, Smart T, Widmer C, Young CM, Vrijenhoek RC: Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Mar Biol. 2009, 156: 395-405. 10.1007/s00227-008-1091-z.View ArticleGoogle Scholar
- Halanych KM, Feldman RA, Vrijenhoek RC: Molecular evidence that Sclerolinum brattstromi is closely related to vestimentiferans, not frenulate pogonophorans (Siboglinidae, Annelida). Biol Bull. 2001, 201: 65-75. 10.2307/1543527.View ArticlePubMedGoogle Scholar
- Rouse G: A cladistic analysis of Siboglinidae Caullery, 1914 (Polychaeta, Annelida): formerly the phyla Pogonophora and Vestimentifera. Zool J Linn Soc. 2001, 132: 55-80. 10.1111/j.1096-3642.2001.tb02271.x.View ArticleGoogle Scholar
- Goffredi SK, Orphan VJ, Rouse GW, Jahnke L, Embaye T, Turk K, Lee R, Vrijenhoek RC: Evolutionary innovation: a bone-eating marine symbiosis. Environ Micobiol. 2005, 7: 1369-1378. 10.1111/j.1462-2920.2005.00824.x.Google Scholar
- Vrijenhoek RC, Johnson S, Rouse GW: Bone-eating Osedax females and their "harems" of dwarf males are recruited from a common larval pool. Mol Ecol. 2008, 17: 4535-4544. 10.1111/j.1365-294X.2008.03937.x.View ArticlePubMedGoogle Scholar
- Knowlton N, Weigt LA: New dates and new rates for divergence across the Isthmus of Panama. Proc R Soc Lond B. 1998, 265: 2257-2263. 10.1098/rspb.1998.0568.View ArticleGoogle Scholar
- Chevaldonné P, Jollivet D, Desbruyères D, Lutz RA, Vrijenhoek RC: Sister-species of eastern Pacific hydrothermal-vent worms (Ampharetidae, Alvinelidae, Vestimentifera) provide new mitochondrial clock calibration. Cah Biol Mar. 2002, 43: 367-370.Google Scholar
- Maas PAY, O'Mullan GD, Lutz RA, Vrijenhoek RC: Genetic and morphometric characterization of mussels (Bivalvia: Mytilidae) from Mid-Atlantic hydrothermal vents. Biol Bull. 1999, 196: 265-272. 10.2307/1542951.View ArticlePubMedGoogle Scholar
- Johnson SB, Young CR, Jones WJ, Warén A, Vrijenhoek RC: Migration, isolation, and speciation of hydrothermal vent limpets (Gastropoda; Lepetodrilidae) across the Blanco Transform Fault. Biol Bull. 2006, 210: 140-157. 10.2307/4134603.View ArticlePubMedGoogle Scholar
- Peek A, Gustafson R, Lutz R, Vrijenhoek R: Evolutionary relationships of deep-sea hydrothermal vent and cold-water seep clams (Bivalvia: Vesicomyidae): Results from the mitochondrial cytochrome oxidase subunit I. Mar Biol. 1997, 130: 151-161. 10.1007/s002270050234.View ArticleGoogle Scholar
- Won Y-J, Young CR, Lutz RA, Vrijenhoek RC: Dispersal barriers and isolation among deep-sea mussel populations (Mytilidae: Bathymodiolus) from eastern Pacific hydrothermal vents. Mol Ecol. 2003, 12: 169-184. 10.1046/j.1365-294X.2003.01726.x.View ArticlePubMedGoogle Scholar
- Hurtado LA, Lutz RA, Vrijenhoek RC: Distinct patterns of genetic differentiation among annelids of eastern Pacific hydrothermal vents. Mol Ecol. 2004, 13: 2603-2615. 10.1111/j.1365-294X.2004.02287.x.View ArticlePubMedGoogle Scholar
- Young CR, Fujio S, Vrijenhoek RC: Directional dispersal between mid-ocean ridges: deep-ocean circulation and gene flow in Ridgeia piscesae. Mol Ecol. 2008, 17: 1718-1731. 10.1111/j.1365-294X.2008.03609.x.View ArticlePubMedGoogle Scholar
- Guinot D, Hurtado LA: Two new species of hydrothermal vent crabs of the genus Bythograea from the southern East Pacific Rise and from the Galapagos Rift (Crustacea Decapoda Brachyura Bythograeidae). C R Biol. 2003, 326: 423-439. 10.1016/S1631-0691(03)00126-4.View ArticlePubMedGoogle Scholar
- Guinot D, Hurtado LA, Vrijenhoek RC: New genus and species of brachyuran crab from the southern East Pacific Rise (Crustacea Decapoda Brachyura Bythograeidae). C R Biol. 2002, 325: 1119-1128. 10.1016/S1631-0691(02)01520-2.View ArticleGoogle Scholar
- Stewart DT, Kenchington ER, Singh RK, Zouros E: Degree of selective constraint as an explanation of the different rates of evolution of gender-specific mitochondrial DNA lineages in the mussel mytilus. Genetics. 1996, 143: 1349-1357.PubMed CentralPubMedGoogle Scholar
- Audzijonyte A, Väinölä R: Phylogeographic analyses of a circumarctic coastal and a boreal lacustrine mysid crustacean, and evidence of fast post-glacial mtDNA rates. Mol Ecol. 2006, 15: 3287-3301. 10.1111/j.1365-294X.2006.02998.x.View ArticlePubMedGoogle Scholar
- Waters JM, Rowe DL, Apte S, King TM, Wallis GP, Anderson L, Norris RJ, Craw D, Burridge CP: Geological dates and molecular rates: rapid divergence of rivers and their biotas. Sys Biol. 2007, 56: 271-282. 10.1080/10635150701313855.View ArticleGoogle Scholar
- Burton RS: Intraspecific phylogeography across the Point Conception biogeogrphic boundary. Evolution. 1998, 52: 734-745. 10.2307/2411268.View ArticleGoogle Scholar
- Hebert PDN, Cywinska A, Ball SL, deWaard JR: Biological identifications through DNA barcodes. Proc Biol Sci. 2003, 270: 313-321. 10.1098/rspb.2002.2218.PubMed CentralView ArticlePubMedGoogle Scholar
- Vogler AP, Monaghan MT: Recent advances in DNA taxonomy. Journal of Zoological Systematics and Evolutionary Research. 2007, 45: 1-10. 10.1111/j.1439-0469.2006.00384.x.View ArticleGoogle Scholar
- Hey J: On the failure of modern species concepts. Trends Ecol Evol. 2006, 21: 447-450. 10.1016/j.tree.2006.05.011.View ArticlePubMedGoogle Scholar
- Mayden RL: A hierarchy of species concepts: the denouement in the saga of the species problem. Species: The Units of Biodiversity. Edited by: Claridge MF, Dawah AH, Wilson MR. 1997, Chapman & Hall, 381-424.Google Scholar
- Avise JC, Wollenberg K: Phylogenies and the origin of species. Proc Natl Acad Sci USA. 1997, 94: 7748-7755. 10.1073/pnas.94.15.7748.PubMed CentralView ArticlePubMedGoogle Scholar
- Jones WJ, Won YJ, Maas PAY, Smith PJ, Lutz RA, Vrijenhoek RC: Evolution of habitat use by deep-sea mussels. Mar Biol. 2006, 148: 841-851. 10.1007/s00227-005-0115-1.View ArticleGoogle Scholar
- Bickford D, Lohman DJ, Sohdi NS, Ng PKL, Meier R, Winker K, Ingram KK, Das I: Cryptic species as a window on diversity and conservation. Trends Ecol Evol. 2007, 22: 148-155. 10.1016/j.tree.2006.11.004.View ArticlePubMedGoogle Scholar
- Ratnasingham S, Hebert PDN: BOLD: the barcode of life data system. Mol Ecol Notes. 2007, 7: 355-364. 10.1111/j.1471-8286.2007.01678.x. [http://www.barcodinglife.org]PubMed CentralView ArticlePubMedGoogle Scholar
- Ivanov AV: Pogonophora. 1963, London: Academic PressGoogle Scholar
- Vrijenhoek RC, Collins PC, Van Dover CL: Bone-eating marine worms: habitat specialists or generalists?. Proc R Soc Lond B. 2008, 275: 1963-1964. 10.1098/rspb.2008.0350.View ArticleGoogle Scholar
- Thewissen JGM, Williams EM: The early radiations of Cetacea (Mammalia): Evolutionary pattern and developmental correlations. Annu Rev Ecol Syst. 2002, 33: 73-90. 10.1146/annurev.ecolsys.33.020602.095426.View ArticleGoogle Scholar
- Deméré TA, McGowen MR, Berta A, Gatesy J: Morphological and molecular evidence for a stepwise evolutionary transition from teeth to baleen in mysticete whales. Sys Biol. 2008, 57: 15-37. 10.1080/10635150701884632.View ArticleGoogle Scholar
- Lingham-Soliar T: Anatomy and functional morphology of the largest marine reptile known, Mosasaurus hoffmanni (Mosasauridae, Reptilia) from the Upper Cretaceous, Upper Maastrichtian of the Netherlands. Philos Trans Biol Sci. 1995, 155-180. 10.1098/rstb.1995.0019.Google Scholar
- O'Keefe F, Carrano M: Correlated trends in the evolution of the plesiosaur locomotor system. Paleobiology. 2005, 31: 656-675.View ArticleGoogle Scholar
- Kear B, Lee M: A primitive protostegid from Australia and early sea turtle evolution. Biol Lett. 2006, 2: 116-119. 10.1098/rsbl.2005.0406.PubMed CentralView ArticlePubMedGoogle Scholar
- Kriwet J, Benton M: Neoselachian (Chondrichthyes, Elasmobranchii) diversity across the Cretaceous-Tertiary boundary. Palaeogeogr Palaeoclimatol Palaeoecol. 2004, 214: 181-194.View ArticleGoogle Scholar
- Kaim A, Kobayashi Y, Echizenya H, Jenkins RG, Tanabe K: Chemosynthesis-based associations on Cretaceous plesiosaurid carcasses. Acta Palaeontol Pol. 2008, 53: 97-104. 10.4202/app.2008.0106.View ArticleGoogle Scholar
- Bardet N: Extinction events among Mesozoic marine reptiles. Histor Biol. 1994, 7: 313-324. 10.1080/10292389409380462.View ArticleGoogle Scholar
- Barbosa J, Kellner A, Viana M: New dyrosaurid crocodylomorph and evidences for faunal turnover at the K-P transition in Brazil. Proc R Soc Lond B. 2008, 275: 1385-10.1098/rspb.2008.0110.View ArticleGoogle Scholar
- Friedman M: Ecomorphological selectivity among marine teleost fishes during the end-Cretaceous extinction. Proc Natl Acad Sci USA. 2009, 106: 5218-10.1073/pnas.0808468106.PubMed CentralView ArticlePubMedGoogle Scholar
- Halanych K: Molecular phylogeny of siboglinid annelids (a.k.a. pogonophorans): a review. Hydrobiologia. 2005, 535-536: 297-307. 10.1007/s10750-004-1437-6.View ArticleGoogle Scholar
- McMullin E, Hourdez S, Schaeffer SW, Fisher CR: Phylogeny and biogeography of deep sea vestimentiferans and their bacterial symbionts. Symbiosis. 2003, 34: 1-41.Google Scholar
- Nelson K, Fisher C: Absence of cospeciation in deep-sea vestimentiferan tube worms and their bacterial endosymbionts. Symbiosis. 2000, 28: 1-15.Google Scholar
- Palumbi SR: Nucleic acids II: The polymerase chain reaction. Molecular Systematics. Edited by: Hillis DM, Moritz C, Mable BK. 1996, Sunderland, Massachusetts: Sinauer Associates, 205-247. 2Google Scholar
- Lenaers G, Maroteaux L, Michot B, Herzog M: Dinoflagellates in evolution. A molecular phylogenetic analysis of large subunit ribosomal RNA. J Mol Evol. 1989, 29: 40-51. 10.1007/BF02106180.View ArticlePubMedGoogle Scholar
- Hillis DM, Dixon MT: Ribosomal DNA: molecular evolution and phylogenetic inference. Quart Rev Biol. 1991, 66: 411-453. 10.1086/417338.View ArticlePubMedGoogle Scholar
- Colgan D, Ponder W, Eggler P: Gastropod evolutionary rates and phylogenetic relationships assessed using partial 28S rDNA and histone H3 sequences. Zool Scripta. 2000, 29: 29-63. 10.1046/j.1463-6409.2000.00021.x.View ArticleGoogle Scholar
- Edgar RC: MUSCLE: a multiple sequence alignment method with reduced time and space complexity. BMC Bioinform. 2004, 5: 113-10.1186/1471-2105-5-113.View ArticleGoogle Scholar
- Maddison WP, Maddison DR: MacClade (Ver. 4.0): Computer Program for Phylogenetic Analysis, version 4.08. 2005, Sinauer Associates, IncGoogle Scholar
- Nylander JAA: MrModeltest, version 2. 2004, Distributed by the author, Evolutionary Biology Centre, Uppsala UniversityGoogle Scholar
- Akaike H: A new look at the statistical model identification. IEEE Trans Autom Contr. 1974, 19: 716-723. 10.1109/TAC.1974.1100705.View ArticleGoogle Scholar
- Rozas J, Sánchez-DelBarrio JC, Messeguer X, Rozas R: DnaSP, DNA polymorphism analyses by the coalescent and other methods. Bioinformatics. 2003, 19: 2496-2497. 10.1093/bioinformatics/btg359.View ArticlePubMedGoogle Scholar
- Xia X, Xie Z: DAMBE: Software Package for Data Analysis in Molecular Biology and Evolution. J Hered. 2001, 92: 371-373. 10.1093/jhered/92.4.371.View ArticlePubMedGoogle Scholar
- Huelsenbeck JP, Ronquist F: MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics. 2001, 17: 754-755. 10.1093/bioinformatics/17.8.754.View ArticlePubMedGoogle Scholar
- Ronquist F, Huelsenbeck JP: MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics. 2003, 19: 1572-1574. 10.1093/bioinformatics/btg180.View ArticlePubMedGoogle Scholar
- Nylander JAA, Wilgenbusch JC, Warren DL, Swofford DL: AWTY (are we there yet?): a system for graphical exploration of MCMC convergence in Bayesian phylogenetics. Bioinformatics. 2008, 24: 581-583. 10.1093/bioinformatics/btm388.View ArticlePubMedGoogle Scholar
- Rambaut A, Drummond A: FigTree: Tree Figure Drawing Tool, version 1.1.2. 2003, Institute of Evolutionary Biology, University of EdinburghGoogle Scholar
- Swofford DL: PAUP*: Phylogenetic Analysis Using Parsimony (*and other methods), version 4.0, Sinauer. 2002Google Scholar
- Drummond AJ, Rambaut A: BEAST, Bayesian evolutionary analysis by sampling trees. BMC Evol Biol. 2007, 7: 214-10.1186/1471-2148-7-214.PubMed CentralView ArticlePubMedGoogle Scholar
- Rambaut A, Drummond AJ: Tracer, version 1.4. 2003, University of OxfordGoogle Scholar
This article is published under license to BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.


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Cyclic generator of a cyclic expression
Dear friends, Given a cyclic expression say in x,y,z like x^4 + y^4 + z^4 - y^2 z^2 - z^2 x^2 - x^2 y^2 what is your more or less elegant way to get a cyclic generator, like x^4 - y^2 z^2 ? Thank you very much. -- --- Francisco Javier Garc=EDa Capit=E1n http://garciacapitan.99on.com


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Music Business Trends and Strategies
Author: Mike King | Course Code: OMBUS-240
This course provides an in depth look at the main structures of the music business, the important changes that are guiding its future, and strategies for creating successful business models. Music Business Trends and Strategies will guide students through the critical areas of the music industry, including publishing and licensing; marketing, promotion, and retail best practices; proper utilization of free music; fan funding and other forms of creative revenue for musicians; current practices for utilizing video, and new business opportunities. By the end of this course, students will have a deep understanding of the pressing issues that all musicians, music industry entrepreneurs, managers, and other music business professionals face in these main segments of the music industry, and how to leverage opportunities that the new business provides.
By the end of this course, you will learn:
- The impact of technology on record label operations and finances
- Current and developing approaches to copyright
- The structure, the traditional and emerging partners, and the revenue opportunities available from music publishing
- Marketing ideas and techniques for building up awareness targeted at your specific community
- Players, payment, and pricing models associated with traditional music retail and distribution outlets
- Nuances and best practices associated with the emerging retail and distribution models
- Approaches for utilizing free music
- Best practices and opportunities available with third party video services
- How musicians are making money, and how to properly prepare and execute on a fan-funded campaign
- New and evolving technological developments that are driving the overall music industry
- Innovative approaches to building a music-focused business
Lesson 1: Inside the Record Business
- Record Industry Overview
- Major Label Consolidation
- The Major Label Approach to Working with Artists
- Recoupable vs. Non-Recoupable Expenses
- How Major Labels are Structured
- Interview with Donald Passman on 360 Deals
- Other Revenue Options for Record Companies
- Alternative Distribution Options with a Major Label
- The Indie Label Approach to Working with Artists
- What an Indie Label Can Do for Artists
- State of the Record Industry
- The Causes of the Decline in the Record Industry
- Roadmap for Record Labels
Lesson 2: Copyright
- Copyright Fundamentals
- The Six Exclusive Copyrights
- Problems with Traditional Copyright
- New Approaches to Copyright New Copy
- Creative Commons
- Cory Doctorow and Creative Commons
- Fair Use
- Psy's Subtle Approach to Copyright
Lesson 3: Publishing
- Music Publishing Overview
- Major and Independent Music Publishers
- Typical Publishing Deals
- Types of Licenses and Revenue Sources for Publishers
- Mechanical License
- Synchronization (Sync) License
- Print License
- Performance License
- The Role of Performing Rights Organizations
- Sound
- Exchange
- Performance Royalty Edge Cases
Lesson 4: Marketing
- New Approaches to Music Marketing
- The Stages of the Internet
- Niche Marketing
- Demographic and Psychographic Overview
- Geographic Data
- Psychographic Data
- Examples of Psychographic Marketing in Action
- Alessandro Cortini and Providing Psychographic-Focused Products
- Jónsi and Alex and Psychographic Collateral Marketing
- Building a Foundation: Acquisition some similar copy to FOM
- Primer for Musicians on Building Up Their Fanbase
- Web Optimization on Your Own Site
- Web Optimization on Third-Party Sites/Search
- Social
- Public Relations
- Touring
- Making it Viral: The Science of Sharing
Lesson 5: Traditional Music Retail and Distribution
- The Foundations of Traditional Physical Distribution and Retail
- Notes from Industry Experts
- How Traditional Physical Distribution and Retail Works
- What a Physical Distributor Does
- The Finances of a Physical Distribution Deal
- Alternative Options for Developing Artists at Physical Retail
- The Putamayo Approach to Retail
- Diversification and Specialization in Physical Music Retail
- Online Distribution and Retail: Structure and Process
- Online Retail Model-Label Involvement
- Online Retail Model-Independent Model
- Online Distribution Overview
- Digital Retail Outlets
Lesson 6: Developing Retail Outlets
- Overview of Digital Recorded Music Retail Models
- The Permanent Digital Download Model
- The Subscription Streaming Model
- The Advertising Model
- The Variable Pricing Model
- The Subscription Streaming Model
- The Past, Present, and Future of the Subscription Model
- Steve Jobs on the Subscription ModelThe Subscription Streaming Payments
- Payments to Rights Holders for Non-Interactive Streaming Services
- Payments to Rights Holders for Interactive Streaming Services
- The Artist's Opinion: Streaming Vs Ownership
- The Utility Model
- Consumer and Artist Perspective
- Bob Lefsetz and Seth Godin on the Utility Model
- "Vaulternative" Retail Model
Lesson 7: Utilizing Free
- Introduction
- The Foundations of Free: Napster, Labels, and Where It All Began
- The Debate Continues
- The "Correlation Does Not Imply Causation" Principle
- Anatomy of a Modern Day Pirate
- How to Utilize Free Music
- The Serialized Approach to Utilizing Free Music
- Noise
- Trade
- Topspin
- Case Study: Billy Van
- Case Study: Berklee College of Music
- Case Study: Wilco
Lesson 8: Video Opportunities and Strategies
- An Overview of Video for Musicians
- Current Video Options for Musicians
- The Social Component of Video
- Case Study: Rufus Wainwright & a Participatory Approach to Social
- Third-Party Video Options for Musicians
- YouTube Live Streaming
- Interview with Evan Lowenstein – StageIt
- Live Video Streaming Case Study: Erin McKeown and her Cabin Fever Series
- Additional Ways to Raise Your Visibility Using Video
- Video Optimization: Musician Case Studies
- Interviews: Karmin's Approach to YouTube, Nicki Bluhm and the Gramblers Van Sessions
- Data and Tracking Your Video's Success
Lesson 9: The Future of Music Coalition
- About the Future of Music Coalition
- 42 Revenue Streams Project
- About the Artist Revenue Streams Project
- Population of Study/Who was Surveyed?
- How Was the Data Collected?
- Research Questions/What was Asked?
- Artist Revenue Streams Project Findings
- Role and Revenue
- The Impact of Technology on Musicians' Earning Capabilities
- Reports
- Policy Issues for Musicians
Lesson 10: Funding your Project
- The Traditional Approach to Funding
- Funding From a Label
- The Traditional DIY Independent Artist Approach to Funding
- Grants
- Sponsorships
- New Funding Opportunities
- Erin McKeown on Her Fan Funding Campaign
- Amanda Palmer: The Art of Asking Ted Talk
- Fan Funding Service Providers: Kickstarter, PledgeMusic, Indiegogo
- PledgeMusic Case Study: Rachael Yamagata
Lesson 11: Technology in Music: Music Discovery, Music Marketing, Music Consumption
- History of Technology in Music
- Facebook, Sharing, and the Social / Open Graph
- Optimization of Streaming Services
- A Three Point Plan for Using Streaming Technologies for Good
- The Freemium Approach to Building Awareness
- Streaming Services: Playlists, Social, and Best Marketing Practices
- Hardware Integration of Streaming Services
- Interview with Peter Gotcher on the Streaming Approach to Marketing Acquisition
- Future of Music Discovery: Filters and Curation
- Pandora
- Nolan Gasser on Creating the Music Genome Project
- Drip.FM
- The Echo Nest
Lesson 12: Music Entrepreneurship and New Business Opportunities
- View from the Top: Roger Brown and Peter Gotcher
- View from the Top: Liz Leahy
- View from the Top: Peter Alhadeff
- View from the Top: Tony VanVeen
- The Lean Startup Approach to Music Business
- Part 1: Vision, True North, Validated Learning, and the MVP
- Part 2: Steer-Test, Measure, Pivot
- Innovative Companies in the Music Business
- Eyes and Ears
- Mobile Roadie
- DigSin
- PledgeMusic
- Nettwerk
Mike King
Author & Instructor
Mike King is a course author, instructor, and the Assistant Vice President for Marketing and Recruitment / Chief Marketing Officer at Berklee Online, the online continuing education division of Berklee College of Music. Prior to working at Berklee, Mike was the Marketing/Product Manager at Rykodisc, where he oversaw marketing efforts for label artists including Mickey Hart, Jeb Loy Nichols, Morphine, Jess Klein, Voices On The Verge, Bill Hicks, The Slip, Pork Tornado (Phish), Kelly Joe Phelps, and Frank Zappa's estate. Mike was the Director of Marketing and Managing Editor of Herb Alpert's online musician's resource, www.artistshousemusic.org, for three years. Mike has written and teaches four courses for Berklee Online: Online Music Marketing with Topspin; Music Marketing 101; Online Music Marketing: Campaign Strategies, Social Media, and Digital Distribution; and Music Business Trends and Strategies. His book, Music Marketing: Press, Promotion, Distribution, and Retail was published by Berklee Press in 2009. Mike was recognized as the Best Music Business Teacher by the National Association of Record Industry Professionals (NARIP) in 2011. Mike has written for Making Music magazine, International Musician, Hypebot, and American Songwriter, and has been quoted in NPR Morning Edition, The Huffington Post, Billboard, The Boston Globe, Wired, CNN, The Boston Phoenix, The Chicago Tribune, Music Connection, and Muso. He's also presented at MIDEM, CMJ, SXSW, NAMM, NARM, SF Music Tech, Futures of Entertainment @ MIT, and Music 2.0. Check out his Website, Blog and follow on Twitter:@atomzooey
Benji Rogers
Benji Rogers is an independent musician from London who has been making his own records since 1999. In 2009, Benji founded PledgeMusic, the leading international direct-to-fan platform offering artists a unique way to engage their fans in the music making process while interacting with a global audience of music super fans, resulting in chart topping albums worldwide. In 2013, Benji was recognized on Billboard's 40 Under 40 Power Players list and in 2014 at the MUSEXPO International Music Awards, he won Digital Executive of the Year.
Jay Coyle
Jay Coyle is the founder and "Music Geek" at the digital marketing firm, Music Geek Services. His company provides artist services for the music industry and has been a Topspin-certified marketer since 2009. Built upon a life-long omnivorous obsession with marketing bands and mixing in equal parts of knowledge gained from an advertising-focused Journalism degree at the University of Georgia along with his skills running a Marketing consultancy for 5 years in Atlanta, Music Geek Services was finally created in 2008 immediately after Jay left a marketing position at the EMI CMG label in Nashville. Jay's first project under the Music Geek moniker was doing work as band archivist and label product manager for Canadian music veterans, Barenaked Ladies. Since then, Jay has worn a lot of hats including sales, marketing, artist development, business development, tour management, and product management. Jay is currently working with Sloan, the Presidents of the United States of America, Jars of Clay, and Carbon Leaf. Jay's core focus is to help further the careers of artists while partnering with them in a "D.I.Y. +" sort of way to have long-lasting and fan-focused careers. To learn more about Jay Coyle or Music Geek Services, visit: Music Geek Services: http://musicgeekservices.com/
Seth Hochman
Seth Hochman is a music business veteran who has worked for Epic Records, Warner Music Group, and Universal Music Group. He has held various marketing and digital account roles working with innovative commercial partners. His most recent role was with Universal Music Group as a Commercial Services Account Director, managing day-to-day business relationships with streaming partners. He launched & marketed UMG's content on Spotify, from it's introduction in the US in 2011 through it's explosive growth stages. As account lead, Seth led UMG's 20+ labels through an evolution in music consumption and revenue growth. Best practices that were developed over three years with Spotify were then focused on video streaming with Vevo, the #1 premium music video channel on YouTube. He worked with labels, artists, and digital marketing teams to develop and monetize visual content. With both Spotify and Vevo, Seth was responsible for Universal's digital advertising strategies which increased download sales and streaming revenue for the company. Prior to working for major labels he was part of the original marketing team that launched Barnes & Noble.com and developed promotional strategies for Kozmo.com, an early forerunner in the on-demand delivery business.
None required
Required Textbooks
None required
Software Requirements
Mac Users
- OS X 10.9 Mavericks or higher (click here for system requirements)
- Latest version of Chrome (recommended), Firefox, or Safari
Windows Users
- Windows 7 or higher (click here for system requirements)
- Latest version of Chrome (recommended), Firefox, or Edge
Hardware Requirements
- 2 GB RAM (4 GB recommended)
- 500 MB hard drive space
- Speakers or headphones
- Webcam
- Internet connection with at least 4 Mbps download speed (http://www.speedtest.net to verify or download the Speedtest by Ookla app from your mobile app store)
Next Term Starts April 3
Duration12 weeks
3-Credit Tuition$1,479
Non-Credit Tuition$1,229


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11Enjoying Life Dont Have ToBe Expencive
Size15 m²
Floor No.0
- Internet/WiFi
- Balcony/terrace
- Lockable room
- Towels
- Kitchen
Accommodation summary
Address | Evesham Road, E15 4AL London, United Kingdom |
Property type | Private room |
Property ID | NYTLFSCO |
Acceptance rate | < 50% |
Response time | A few hours |
Calendar updated | 2 days ago |
The guest have access to the kitchen and bathroom, washing facilities, towels and bedding, small rear garden and free wifi.
-We are located in STRATFORD, EAST LONDON, TRAVEL ZONE 3
- we are 7 minutes away from the London city center (from Liverpool Street)
- we are 10 mins away from Stratford underground station by walking
- we are 25 mins away from Oxford Street
- we offer good price and quality adjusted to this price
- we taking needs of our guests seriously :-)))
Stratford area is a classic example of East London marked by incredible mixture of cultures, races, traditions. It is the heart of Olympic Games in the nearest past.
This area exceptionally well connected with city center literally 7 minutes away from Liverpool Street by train.
This area is also very convenient for travelers from/to STANSTED AIRPORT thus many of the coaches go to London through STRATFORD. However other airports are also fine due to well-developed underground, train and bus connections in the city.
All TRAVEL CARDS and OYSTER CARDS are accepted. What you need is an Oyster card with One Day Travel Card Zone 1-3 cost approximately £5.70 per day. London is an expensive town so be prepared to spend.
Take Central Line underground train to Stratford. From Stratford the house is withing walking distance. (website hidden) the address previously and take the map with you.
We are 7 minutes away from Liverpool Street station by train or underground. Liverpool Street is a perfect spot to start exploring London. Liverpool Street is in the heart of London's business City- with its panoramic views, dramatic St. Paul Cathedral, well known Tower Bridge, Tower of London and River Thames itself. It is highly recommend for every tourist to use the Liverpool Street / Stratford route for his/her trips thus it will save lots of time, energy and frustration. To remind you again: WE ARE 7 MINUTES AWAY FROM the LIVERPOOL STREET STATION BY TRAIN.
Please use this tip to help yourself make your trip to London as pleasant as possible :-)
House rules
Details |
Maximum nights | 1 month |
Check-in after | Noon |
Check-in before | On arrangement |
Check-out before | 11 AM |
Cancellation Policy |
Reviews for this host's other properties
Simple house, nothing amazing but the essential thing that is well-connected to the center that stantded since the house is located a few stops from Stratford where we find the bus to the airport and vice versa (terrevision)
Casa semplice, niente di stupendo ma la cosa essenziale che é ben collegata sia con il centro che a stantded dato che la casa si trova a poche fermate da stratford dove troviamo il bus che porta all'aeroporto e viceversa(terrevision)
We are satisfied about our stay in this room. Simple room for a good price. Close to the Underground station Upton Park and easy to find, thanks to the good information of the host.( we did not see him, but this was no problem) Supermarkets are also very close.The house was very quiet, with friendly other people. A next time we would like to come back here.
That is not Hilton but you get all you need. Big benefit for me is there is West Ham staduim very close.
We booked a room with this host in January to travel in April. Ruslans kindly provided us to answer questions about transport etc, but when the day our flight was canceled by air traffic controllers strike and could not go, this being no fault of us and to be the STRICT cancellation, but we had no right to return of 5%. We got in touch with Ruslans to reach an agreement but gave no indication until the day after.
We got in touch with Windu and after contact for an agreement Ruslans his answer was that he could not refund our money (or rather%) because I had to cover the costs of that day.
After contacting the next day, I do not answer to our suggestion back somewhat% What we said was that if I ever came back would make us a good deal.
In short, strict cancellation is STRICT 100% no matter the cause.
Reservamos una habitación con este anfitrión en enero para viajar en abril. amablemente Ruslans siempre nos contesto a las dudas sobre transporte etc, pero llegado el día nuestro vuelo se cancelo por huelga de controladores aéreos y no pudimos ir, siendo estas causas ajenas a nosotras y al ser la cancelación ESTRICTA, no teníamos derecho más que a la devolución del 5%. Nos pusimos en contacto con Ruslans para llegar a un acuerdo pero no dio señales hasta el día después.
Nos pusimos en contacto con Windu y tras ponerse en contacto con Ruslans para un acuerdo su respuesta fue que no podia devolvernos el dinero ( ni un tanto %) porque tenia que cubrir los gastos de ese día.
Tras contactar con el el día siguiente, no contesto a nuestra sugerencia de devolución de un tanto % Lo que nos dijo fue que si volvíamos algún día nos haría una buena oferta.
En resumen, la cancelación ESTRICTA, es ESTRICTA 100% da igual las causas.
First, the positive: very low, kitchenette and washing machine. In addition, a towel and bedding is provided each. In addition, very easy on and off.
What bothered us: very poor condition of the house, very noisy. The sanitation facilities are in very poor condition, partly cracked, defective.
If you want to stay there should bring toilet paper, disinfectant and a sense of smell insensitive.
Zuerst das positive: sehr günstig, Küchenzeile und Waschmaschine vorhanden. Ausserdem wird je ein Handtuch und Bettzeug gestellt. Zudem sehr einfaches ein- und ausziehen.
Was uns gestört hat: sehr schlechter Zustand des Hauses, sehr hellhörig. Die sanitären Anlagen sind in sehr schlechtem Zustand, teilweise gesprungen, defekt.
Wer dort übernachten möchte sollte sich Toilettenpapier, Desinfektionsmittel und einen unempfindlichen Geruchssinn mitbringen.
Good value for money. Well located, close to the subway, shops and bars! :)
Buena relación calidad precio. Bien situado, cerca del metro, tiendas y bares! :)
It was good time in London! Nice, warm, cozy room (if it was not enough warm we have small electric radiator), clean kitchen and bathroon in a typical english residential building. We could without problems prepare something to eat. Householder was helpful and nice. We saw him once at the beginning.
- Rooms: relatively small, bed very unstable (is on wheels and is more tinkered together than anything else), the walls are very thin so you can hear every conversation throughout the house and all other activities in the house.
- Location: 15 min walk to the train and from there no 40 min to downtown
- WC: As already described in other reviews has occurred, the lack of toilet paper problem with us.
-Kitchen: Everything there that you need :)
- Zimmer: relativ klein, Bett sehr instabil ( steht auf Rollen und ist eher zusammen gebastelt als alles andere), Die Wände sind sehr dünn so das man jedes Gespräch im ganzen Haus sowie alle anderen Aktivitäten im Haus hört.
- Lage: 15 min Fußweg zur Bahn und von da keine 40 min in die Innenstadt
- WC: Wie schon in anderen Bewertungen beschrieben ist das mangelnde Toilettenpapier Problem auch bei uns aufgetreten.
-Küche: alles da was man braucht :)
Everything was excellent.
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text:
Moderators: Jenise, Robin Garr, David M. Bueker
Larry Greenly wrote:Do you eat dessert before or during the main course? And what are your thoughts concerning this? (And my apologies if you were the woman sitting next to me.)
Larry Greenly wrote:And my apologies if you were the woman sitting next to me
Howie Hart wrote: The first is I eat slow. I used to eat fast (a habit acquired while in the Marines), but when my wife and I were dating, she would eat slow, saying it a technique to curb the amount of food one eats.
I want 'em together, dammit!
Larry, as you know, there is a widespread philosophy about eating dessert first, because you don't know for sure if something won't come up before you get to it otherwise.
Stuart Yaniger wrote:if you try to get that combination at the end of a dinner in France or Italy, they'll either spit in your food or ignore your request altogether.
FLDG Dishwasher
Tue Mar 21, 2006 3:45 pm
The Pacific Northest Westest
Karen/NoCA wrote:My career was working with seniors who volunteered with our City. I'm guessing that the lady who ate her dessert alternately with her meal, was loving the fact that she actually had dessert.
Many seniors who live alone do not eat properly and do not take the time to cook anything that requires a bank of time and work. You will notice that at many senior events the table is set with the dessert plated. The hot food is usually buffet style or served after everyone is seated. It was not unusual in the least for me to see the gals (and guys) picking away at dessert before the food even came out.
Karen/NoCA wrote:Many seniors who live alone do not eat properly and do not take the time to cook anything that requires a bank of time and work.
Karen/NoCA wrote:I wasn't applying any rule to seniors. I was replying to Larry's post about an obervation he made with a senior group. I responded with observations I had made in my 16 years of working with seniors. I've seen children doing the same thing.
Golfball Gourmet
Thu Oct 12, 2006 9:12 am
Extreme Southwest & Extreme Northeast
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text:
Polk County Prosecuting Attorney Kenneth Ashlock filed a motion Monday for a mental exam for Blaec Lammers, 20, Bolivar, who is charged with plotting a shooting spree at a showing of the final chapter in the Twilight series, "Breaking Dawn."
Lammers is being held in the Polk County Jail in lieu of $500,000 bond.
Online court records show Polk County officials have charged Lammers with assault in the first degree, making a terroristic threat, and armed criminal action.
No court date has been set.


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Woodburn Wooden Shoe Tulip Festival
Tiia Vissak 4/17/2012 8:57:14 AM
great colors, details & dof! #1457108Liz blahh 4/17/2012 9:45:21 AM
This is really beautiful,like the tractor in the midst of all that colors! #10095455Susan M. Campbell 4/17/2012 11:18:33 AM
I bet this was awesome to see in person. I love your selective focus on this. A great capture. It kind of reminds me of our flower fields pictures. #10095564Carol Quina 4/17/2012 12:19:55 PM
Love it! So PRETTY! #10095640Jill Odice 4/17/2012 12:38:38 PM
LOve your version of the truck ! What a great place to visit! Thanks! #10095664Mike Clime 4/17/2012 2:12:23 PM
Beautiful capture and colors and I'm a big fan of old JDs. #10095789Terry Cervi 4/17/2012 4:06:13 PM
Wow, look at all of those beautiful tulips! What a great shot, John! #10096008Darren K. Fisher 4/17/2012 4:13:33 PM
I would do anything to see something like this in person. Your capture and really beautifully done. #10096033Shelly A. Van Camp 4/17/2012 6:33:35 PM
Beautiful....come here Darren. The Botanical gardens are full of tulips now. #10096242Linda L. Ruiz 4/17/2012 9:25:58 PM
Tons of comments. @WOW.com. This came out fantastic!!!!! #10096459Inge Linden 4/18/2012 1:35:32 AM
Wonderful capture and colors! #10096633Corinne Hurt 4/18/2012 9:53:49 AM
Very nice! #10097436Marilyn K. Lincecum 4/18/2012 11:07:41 AM
AWESOME color...wonderful comp, DOF and detail. #10097537Sabrina Ryan 4/30/2012 7:36:31 AM
Beautiful color and wonderful pov, David! #10114793
Exif: F Number: 2.8, Exposure Bias Value: 0.00, ExposureTime: 1/800 seconds, Flash: did not fire, compulsory flash mode, ISO: 100, White balance: Auto white balance, FocalLength: 100.00 mm, Model: Canon EOS 50D
Uploaded on 4/17/2012 8:15:40 AM
Camera: Canon EOS 50D Digital SLR Camera
Copyright for this gallery photo belongs solely to David Hurt. Images may not be copied, downloaded, or used in any way without the expressed, written permission of the photographer.
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text:
Memento Moron: Remember, Thou Art Stupid
Sunday, January 27, 2008
New Quote of a Lifetime
"I be chef like da-da."
- The Lad
three years to the day
, at that. No, dammit, I just got some onion in my eyes from cooking. Shut up.)
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