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Galaxy Note 6 Release Date, Specs, Features And More
Samsung's next flagship is said to be the Galaxy Note 6. Note 5 was released last year along with S6 Edge+. Unfortunately, none of these two devices did well. So, this year after Galaxy S7 and S7 Edge, Samsung may release another Note with advanced features. This device may be a mixture of Note 5 and the latest S7 phones. Get more details about Galaxy Note 6 release date, features and more here:
Samsung's Galaxy Note 6 release date is rumored to be somewhere in August this year. As you know, currently only the latest phones from the company support virtual reality, and now, this upcoming flagship may be another device that supports Gear VR. Any smartphone, no matter which brand it belongs to, cannot be judged without going through the specs list.
According to Android Central, Galaxy Note 6 will sport a 5.8-inch screen and will come with 4K Super AMOLED display. The report also suggests that this phone from Samsung will pack a "battery substantially larger than the 3,000mAh cell of the Note 5."
Speaking of the chipset, it is not clear whether Samsung will use only Snapdragon 820 or like Galaxy S7 and S7 Edge opt for a combination of Exynos and Snapdragon chips. Experts say that the performance of S7 series varies depending on the chips used. While Snapdragon 820 phones are the best, Exynos phones, though good, are not excellent. The chances are that based on these reviews, Samsung may completely drop Exynos chips and choose Qualcomm for the upcoming high-end devices.
Related: Android N Hints At Galaxy Note 6 Specs?
Let us now move on to the Galaxy Note 6 features. The phone, just like Galaxy S7 and S7 Edge, will be water-resistant. It will also support wireless charging and microSD card. This upcoming smartphone from Samsung may be the first device to don a USB Type-C port. Though nothing has been confirmed by the company yet, these rumors cannot be totally denied as most of the S7 and S7 Edge rumors also turned out to be true.


text:
The wrapping of the wrapping paper book
on May 19, 2017 at 8:51 am
Posted In: Upcoming releases
Ah, finally we have something that looks like an actual cover for the upcoming A Charlie Brown Christmas Wrapping Paper Activity Book. The few uncolored figures to the left look to me like ghosts in the composition. Maybe they're the ghosts of Christmases Past, Present, and Future?


text:
What is the best forum plugin?
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text:
Delta County's local government designee (LGD) for oil and gas regulation issues, Bruce Bertram, has retired from the post.
At the Board of County Commissioners' meeting on Monday he received compliments and thanks for his years of service to the county.
Oil/gas designee retires
Delta County's local government designee (LGD) for oil and gas regulation issues, Bruce Bertram, has retired from the post.
Agricultural burn generates complaints
County commission chairman Doug Atchley said on Monday that he has asked the county and state health departments to provide statements on a situation at Eckert where smoke from an allowed open burn operation is impacting residents of Horizons West Mobile Home Park and Horizons Care Center.
POWs/MIAs remembered
Members of Delta's Lee Marts Veterans of Foreign Wars Post and Auxiliary #3571, and members of Delta's American Legion Post #65 conducted the Missing Man Table ceremony at Arby's in Delta last Friday.
They said that ceremonies would also be conducted at the Delta Area Chamber of Commerce office, and at Starvin' Arvin's.
County clerk's office hours change Oct. 2
An earlier closing time for business at the Delta County clerk and recorder's office in the courthouse was approved by the county commissioners on Monday. Beginning Oct. 2, the closing time will be 4:30 p.m.
Orchard City has no jurisdiction in burn complaint
The Orchard City Town Board, at its regular meeting on Sept. 13, heard complaints from residents of Horizons West mobile home park about smoke from burning piles of brush near the retirement community.
Deann Thomas, a resident of the park, and three of her neighbors told the town board of smoke and other problems they claim were caused by the burning of brush just north of their senior mobile home park on Running Deer Road.
Repealing marijuana bans on hold until April election
The Orchard City Town Board acting at its regular monthly meeting on Sept. 13 voted to rescind its previous decision to proceed with repealing current bans on marijuana business in town.
The town board also voted to place three referred measures on the April 2018 ballot asking voters to decide on new revenue measures for town government.
Senior Hikers celebrate their heritage
Mel Edmunds would have totally enjoyed the 15-year reunion of the Cedaredge Hikers at Town Park last week.
After all, having the event was his idea to begin with.
Ag burning complaint set for OC board meet
Deann Thomas, a retired physical therapist, says she will raise concerns with the Orchard City Town Board about an agricultural burn operation next to the senior mobile home park where she and elderly neighbors live.
Orchard City now eyeing 3 tax questions on April ballot
Orchard City's Cannabis Revenue Committee announced last week that it is ready to drop a recommendation for immediate repeal of the town's ordinances which bans marijuana business. The committee had recommended the town repeal current bans on medical and commercial marijuana businesses before taking the question of sales tax from marijuana businesses to the voters. It has drawn notable public opposition.
Events of 9/11 commemorated
A ceremony commemorating the 16th anniversary of the 9/11 attacks was held in Delta Monday, Sept. 11, at Bill Heddles Recreation Center.
The event was organized and conducted by members of Delta's Lee Marts Veterans of Foreign Wars Post and Auxiliary #3571.
Pops in the Park -- 10 years of enriching lives
One of Surface Creek Valley's notable lifestyle enhancements took place last Saturday in Cedaredge Town Park.
The 10th annual Pops in the Park concert by the Valley Symphony Orchestra and Chorus brought a large number of people with chairs, coolers filled with snacks, and their umbrellas for an evening of symphonic music and song.
State pledges more for enhancement project
The Cedaredge Town Board on Sept. 7 agreed to an offer from CDOT to contribute up to $75,000 more toward eventual completion of the Main Street Enhancement Project. Of the total $75,000, the town will be required to contribute 20 percent, or about $15,000, it was stated at the meeting.
An impetus to create
Karen Brueggemann has been creating objects of art her whole life. She has been selling her creative works since she was in junior high school.
But it has been only since she and Ted, her husband of 54 years, began their move to Cedaredge 10 years ago that Karen really began to think of herself as an artist.
Fighting Parkinson's together
A Cedar Mesa couple has taken up the fight against an incurable, debilitating disease and the limiting effects it has on people's lives.
Charlie Farrell and Jan Blue this week bring their personal ministry to the community by making available to Parkinson's disease sufferers at no cost a new therapy regimen. They explain it has shown proven results in helping people overcome symptoms of Parkinson's.
A burning passion
When Scott Sullivan retired from his career at the Delta Correctional Center after 16 and a half years, art became a much bigger part of his life.
With new-found free time he now had for pursuing his passion, he completed construction of a studio behind his house just north of town in Cedaredge.
In her element
Sandy Wilt's near 30-year-long affection for the photographic hobby has filled her life with enjoyment, adventure and many friendships. She looks forward to a future of excitement from her avocation, a future which, perhaps surprisingly, she views as "a little bit scary, too."
Repurposing with flair
An Eckert man has taken his talent for art, merged it with lessons his dad taught him years ago and created a vocation for himself and a distinctive type of craftsmanship.
John Good found early in life that he had a talent and love of art which he pursued into his adult years.
What lies below
The geologic features of Delta County that contribute to its bounty of agriculture and natural resources, and to the scenic beauty and recreational pleasure it affords have been the professional province of Dave Noe of Paonia.
He is a geologist "recently retired" from the Colorado Geological Survey.
Art of giving
Art is part of life, and life is about giving. Cedaredge area artist Ella Kelly has found that art and giving have been center points in her personal and professional life.
"We are so blessed to have people in our lives who teach and help us with their talents," Ella says.
A class act
Music audiences in this area have a new entertainment option to enjoy, a real "Class Act." The group offers a wide range of classic songs to enjoy; a performance style that is at once both engaging and relaxing; and a catalog of numbers presented with a true love of the music they play.
Pickleball, anyone?
The Grand Mesa Picklers is a growing circle of local friends and newcomers who quickly become friends. They share their enthusiasm for quality of life, for physical fitness activity, and for playing together on a regular basis.
A model of success
With local economic development ideas turning once again to themes of rural gentrification, lifestyle marketing, recreation and tourism there is one local economic development asset that provides a model and foundation of success to build on.
It is the Grand Mesa Scenic and Historic Byway.
Creative community
Art can't thrive in isolation -- it needs a community of artists and a community for expression.
A new artists' cooperative gallery in Cedaredge is providing both types of community connection for local artists who have a new venue for display and sale of their work.
A saga of survival & faith
Cedaredge native Keith Loucks has published a novel about adventure and survival -- a book that entertains as it educates, and one that examines deeper questions of living Christian faith in a dangerously fallen world.
In the 405-page novel titled "I Wish I Had An Orange," the normal, everyday American life is reduced in a moment to the level of Third World deprivation when a nuclear device is exploded in the upper stratosphere 30 miles above.
A life turn to art
When Sue Jochum decided it was time to leave the corporate world of high technology and pursue her life-long passion for art, she chose the Surface Creek Valley as home to be near other family who live here.
Now, with her computer programming days behind her, she is finding the time to enjoy plein air painting sessions with friends, to participate with her work in shows and exhibitions, and to develop an enterprise in portrait painting, a discipline that she has discovered a natural talent for.
Champions of Nordic skiing
The Grand Mesa Nordic Council is marking a 25th year of providing great outdoor recreation experiences for local skiers and other winter sports enthusiasts.
Over the course of those 25 years, the GMNC has also contributed significantly to developing a first-class complex of trails for cross-country skiing, snowshoeing and skijoring that is a tourism draw for the county.
Driven to serve gladly
Becoming part of a small community means connecting with people.
There is no surer way for someone to know they're making a positive difference and connecting with the heart of a community than to find an avenue of service.
Drones give the economy a lift
Drones have arrived in Delta County -- but they aren't the spying, harassing kind of drones that are reported on the national newscasts.
A Cedaredge company is helping bring the high-tech surveillance craft here for economic development uses that range from marketing promotion to improving agricultural yields.
Wheelin' 'n dealin'
One event noticeably absent from Delta's annual cycle of summer activities is a big car parts swap meet.
Last Saturday, that gap in the community events calendar was filled in by a well-attended auto parts swap at Confluence Park.
The promise of America
Larry and Dawn White took America at its promise and proved to themselves that the promise of America is good.


text:
Chelsea Home Furniture 75E18020
75E180-20 Ramsey Accent Chair
The Chelsea Home Furniture Ramsey accent chair in roundabout mineral finish richly crafted wood frame and plush fabric brings a sense of Victorian elegance to any living room area. This beautiful set epitomizes Chelsea's legendary reputation for quality and comfort. This collection's elegant royal style and warm colors will extend an open invitation for everyone. Available at AppliancesConnection
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- Brand:Chelsea Home Furniture
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text:
International Security
a highly competitive world of rapid changes
security is becoming a hot topic of growing
importance. Developing efficient security
patterns is a prerequisite for the prosperity
and well-being of nations of limited resources
and technology. Thus, the Institute aims at
initiating and sponsoring specialized research
in the field of security.
The Arab Institute for Security Studies strongly
believes that security is not strictly a
military issue but rather a comprehensive term
which necessarily have to deal with water,
environment, refugees, disarmament among other
issues. Thus, Security is multidisciplinary
issue which should involve the widest spectrum
of individuals and benefit from the contribution
of all sectors in society.
The Center holds an annual inter-regional
conference in Amman that addresses "Security
Concerns in the Middle East", where a
multi-disciplinary group of experts meet to shed
light on various aspects of regional security in
the Middle East in addition to viewing and
comparing security models and arrangements
elsewhere in the world. Outcomes of such
meetings are delivered to authorities and
decision makers in Jordan and the region.
The Institute examines other aspects of security
I- Foreign Policy Analysis
The Institute sponsors a "National Committee for
Foreign Policy". The committee observes
developments on the local, regional and
International scenes and analyses possible
impacts on the region.
II- Environmental Security
NBC security represent one of the Institute
priorities (nuclear, biological and chemical
materials). The Institute is under obligation to
improve practices relating to NBC management at
facilities dealing with such materials
(including other categories of hazardous
materials as well as hazardous waste). Work
includes provision of feasible solutions as well
as developing contingency plans, thus protecting
citizens health and environment. Recently, the
Institute was successful in establishing the
"Commission for monitoring biological and
chemical hazards in Iraq". The Institute possess
technical and monitoring capabilities including
radioactive monitoring facilities and
specialized chemical analysis laboratories.
III- Biosecurity
Biosecurity represent another area of interest
and falls within the mandate of our Institute.
Of particular interest is the status of
biological weapons and the development of
biological warfare capabilities. On the
practical level, the Institute is active in
developing guidelines and procedures for
organizations dealing with biological agents (as
well as chemical agents). Additionally, the
Institute cover other areas pertaining to
disease surveillance, infection control … etc.
IV- Disaster Management and Risk Reduction
The Institute supports the establishment of a
"Regional network for Disaster Management and
Risk Reduction". The Network is a joint
initiative lead by the our Institute in
cooperation with a number of International
agencies. It is entrusted to develop disaster
management capabilities, introduce risk
reduction practices and facilitate the adoption
of new technologies. Additionally, the Network
is engaged in the following tracks:
A- Developing a regional policy on Disaster
Management and Risk Reduction
B- Promoting regional research and developing
country reports on preparedness of countries to
disasters (natural and man made).


text:
Self Storage Crivitz
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|Sandstone Storage
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text:
About this Business
- HoursSunClosedMon-Fri10:00 AM - 6:00 PMSatClosed
From Our Editors
Old Village Medispa offers high-quality and specialized medical treatment for those in the Mount Pleasant area and beyond. Bring your skin back to life with a facial professionally done by this clinic's expert aestheticians. Get healthy, gorgeous skin when you take advantage of a skin care treatment from this clinic, such as acne treatment, chemical peel, IPL, microdermabrasion, and spider vein removal. To reduce development of deep wrinkles, head to this clinic for an injection. When those pesky hairs just won't go away, visit Mount Pleasant's Old Village Medispa for a close hair removal treatment. When you need medical treatment on the fly, this clinic is readily available to assist you. To maintain a timely workflow, this establishment is strictly appointment only. No cash, no problem. This beauty salon happily accepts payments by major credit card. With a parking lot adjacent to Old Village Medispa, you won't get stuck circling the block. The next time you're in need of a doctor, be sure to stop in at Old Village Medispa.


text:
"Open Russia" has elected a new Board headed by Khodorkovsky
The project "Open Russia" the former head of Yukos Mikhail Khodorkovsky held a founding conference, electing the Council, formalizadas structure and planned activities: the movement will support citizens in disputes with the authorities and companies, as well as participation in regional elections, reports "Interfax" with reference to the press service of the movement.
The founding conference of the "Open Russia" is held in the Finnish capital Helsinki. There approved the program, Manifesto and Charter. According to Khodorkovsky, the immediate task of the movement – legal support in disputes with authorities and monopolies, public control over activities of the authorities and the media, as well as participation in elections to municipal councils and the regional legislative Assembly.
The conference also elected 11 members of the Board of the movement. In addition to Khodorkovsky, the Board includes: co-Chairman of Parnassus Vladimir Kara-Murza Jr., head of St. Petersburg branch of the party Andrei Pivovarov, the candidates of the "Open Russia" on last elections in the state Duma Maria Baronova, Olga zhakova, Savin Egor and Marina Belova. None of them to be elected to the Duma failed.
The members of the Council of the movement "Open Russia", elected at the founding conference on November 12. pic.twitter.com/lzE32CBr9a


text: "Mikhail Gorbachev and Daisaku Ikeda are contemporaries raised in different cultures: Gorbachev is a statesman whose origins are the Marx-inspired world of Communism, while Ikeda is a Buddhist inspired by the thirteenth century Japanese sage, Nichiren. Moral Lessons of the Twentieth Century emerged from a series of conversations between these two men. Together they explore their experiences of life amidst the turmoil of the twentieth century and together they search for a common ethical basis for future development. They conclude that values are born of culture and that peace, progress and social justice can only be achieved through sincere communication and cultural exchange. Humanisme--20e siècle 2005 New York Histoire universelle--20e siècle--Philosophie I.B. Tauris Political ethics 9781850439752 1850439753 2005 57484270 Moral lessons of the twentieth century : Gorbachev and Ikeda on Buddhism and Communism 57484270 As the new century begins, they have sought to turn the spotlight on the challenges which face humanity. The book is a call for dialogue in pursuit of values that bridge culture and time."--Jacket. en History, Modern--1989- Morale politique--20e siècle History, Modern London Communisme 2017-09-02 Communisme--20e siècle In the United States of America and in Canada distributed by Palgrave Macmillan Since 1989 Valeurs sociales--20e siècle enk Ikeda Daisaku Daisaku Ikeda Gorbachev Mikhail Sergeevich Mikhail Sergeevich Gorbachev 1931 9781850439769 1850439761 Boeddhisme


text:
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A New Book for the Serious Coin Collector: Inside the Rare Coin MarketplaceIn Bressett?s Whitman Guide to Coin Collecting, a newcomer will learn about coin collecting as a hobby; reasons people collect coins; coins as historical documents; how coins are made; where to find them; what factors affect their value; and the basics ...Good photos of bad coins: An example of crowdsourcing and numismatic publishing... the Whitman Guide to Coin Collecting The 288-page softcover Whitman Guide to Coin Collecting debuted July 4, 2017, and is now available from booksellers and hobby shops nationwide, and online (including at the official website of Whitman Publishing ...Indian Head cent gave hobby boostWhen coin collecting became a national hobby in the 1850s it was due to the withdrawal of the old copper cents and half cents. Ordinary citizens enjoyed finding all of the different dates that could be obtained. In the 1920s and early 1930s numismatics was ...Pembrokeshire metal detectorists unearth hidden treasuresHe was interested in coin collecting but could not afford to buy coins, so his mother suggested he buy a metal detector to find some. The retired plumber from Saundersfoot said detecting had been his "lifelong hobby". "If you find something, your interest ...What Amazing Sports Plays Are No Longer Special?I don?t have time to take up 17th-century coin collecting, man. Sometimes it?s okay to live a generic existence. Also, not everyone is gonna volunteer their hobby on the spot, or even realize that their hobby IS a hobby. I like to cook, but shine a ...Silver Coins Jewellery Price ListMany people collect silver coins as a hobby called numismatics. Coin collecting is a fun and engaging hobby that combines history with leisure. Though silver is a metal that?s inferior to gold, it?s a metal that's worth investing in. Similar to gold ...Cleveland man says coin collecting hobby is time well-spentThe hobby of coin collecting can be traced back as far as the Renaissance, when it became popular among some members of the wealthier class to present old and exotic coins to guests in the midst of festivals and other special occasions. For years, many ...RGV Coin Collecting Becoming a Popular HobbyMCALLEN ? It?s evident in the Rio Grande Valley that coin collecting is the fastest growing hobby. So much, that the Hidalgo Coin Club has had to move to a bigger location for its monthly coin shows. The McAllen Chamber of Commerce hosted a ribbon ...Coin Collecting: Investors? Hobby With High StakesNumismatics: a hobby to some, a gamble to others and a serious investment to many collectors. The aim is the purchase of proof coins for pleasure and also a profit. The players range from small children hopeful of finding a 1913 Liberty head nickel in a ...Winona Coin Club has annual coin show on SundayAround that time is when Schneider picked up the coin collecting hobby himself. He said he was looking through change in pockets, coin jars and couch cushions, hoping he would find some of those pennies. Schneider said he enjoys coin collecting because of ...
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- The Paleontological Society
Bivalve death assemblages from subtidal environments within the tropical Bocas del Toro embayment of Caribbean Panama permit a test of the extent to which levels of damage are determined by the intrinsic nature of shell supply (proportion of epifaunal species, thick shells, calcitic shells, low-organic microstructures), as opposed to the extrinsic postmortem environment that shells experience. Only damage to interior surfaces of shells was used, to ensure that damage was unambiguously postmortem in origin. We find that facies-level differences in patterns of damage (the rank order importance of postmortem encrustation, boring, edge-rounding, fine-scale surface degradation) are overwhelmingly controlled by environmental conditions: in each environment, all subsets of the death assemblage present the same damage profile. The composition of shell supply affects only the intensity of the taphonomic signature (i.e., percentage of shells affected), and only in environments containing hard substrata (patch reefs, Halimeda gravelly sand, mud among patch reefs). In these environments, epifauna, whether aragonitic or calcitic and whether thin or thick, exhibit significantly higher damage than co-occurring infauna, probably due to the initial period of seafloor exposure they typically experience after death. Thick shells (>0.5 mm), regardless of life habit or mineralogy, are damaged more frequently than thin shells, probably because of selective colonization by fouling organisms. Calcitic shells show no consistently greater frequency of damage than aragonitic shells, and high-organic microstructures yield mixed patterns. Taphofacies surveys in such depositional systems could thus be confidently based on any subset of the fauna, including diagenetically residual assemblages of calcitic shells and thick-shelled molds. Further tests are needed to determine whether the higher levels of damage observed on some subsets of shells are a consequence of greater time-averaging (thus lower temporal resolution), greater exposure time, preferential attack (potential bias in relative abundance), or some combination of these. Paleobiologically, however, the implication is that ecological subsets of bivalve assemblages are not isotaphonomic, either in tangible damage or in probable bias, within hard-substrate environments, although they may be within soft-sediment environments. In actualistic studies, targeting broad classes of taxa for comparison across environments maximizes our ability to extrapolate taphonomic guidelines into the fossil record, where life habits, skeletal types, and shallow subtidal habitats have dramatically different patterns of abundance and deployment.
Paleontologists have long recognized that the quality of fossil preservation varies among environments (see historical review by Cadée [1991]). The classic actualistic work on this phenomenon was by Schäfer (1972), who differentiated a series of idealized biofacies on the basis of death assemblage characteristics (are taxa largely indigenous or exotic to the facies? disturbed from life positions? damaged by postmortem processes?). "Taphofacies analysis" (Speyer and Brett 1986) has increasingly focused on the quantitative damage profile or "taphonomic signature" of death assemblages (e.g., observed frequency of bored and encrusted hard parts, extent of skeletal disarticulation and fragmentation), with the aim of developing a set of criteria for paleoenvironmental interpretation and for recognizing between-habitat transport of shells (e.g., Davies et al. 1989; Staff and Powell 1990; Anderson and McBride 1996; Brachert et al. 1998).
One aspect of marine taphofacies that has remained unclear is the extent to which facies-level differences in the quality of preservation are due to differences in postmortem conditions as opposed to differences in the material supplied by living organisms. That is, what is the relative importance of the extrinsic environment of accumulation (destructive physical, chemical, and biological agents, scales of time-averaging), as opposed to the intrinsic attributes of tissues produced or otherwise introduced to that environment (e.g., types of hard-part construction, condition upon death)? Determining the relative contributions of these factors would permit a better evaluation of (1) the limits of extrapolation back into the fossil record, where the composition of skeletal supply differs owing to evolution (e.g., different proportions of epifauna and infauna in soft substrata, of thick versus thin shells, of shell mineralogies and microstructures), and (2) the relative tendencies of different classes of skeletal supply to accrue postmortem damage, and thus how best to target field effort when conducting taphofacies analysis for paleoenvironmental differentiation. The results also bear on the question of (3) differential bias (underrepresentation) of taxa, and particularly of entire subsets of faunas, in death assemblages. Although direct experimentation is necessary to establish causation, strong links between intrinsic attributes and particular levels or patterns of damage indicate the potential for bias in preservation.
Taphofacies analyses have tended to focus on the role of extrinsic postmortem conditions in generating damage profiles, but most workers clearly appreciate that both intrinsic and extrinsic factors (nature and nurture) are important (Meyer et al. 1989). However, distinguishing these effects can be difficult. Nebelsick (1992), for example, attributed much of the variation in damage profiles of echinoid death assemblages in Safaga Bay, northern Red Sea, to the ecological dominance of thin-shelled regular echinoids in hardground habitats and of robust clypeasteroids in sands, but the spatial segregation of taxa was too strong to differentiate taxonomic and environmental effects. Similarly, Parsons and Brett (1991) postulated that differences in shell supply probably affected taphonomic signatures in molluscan death assemblages from the northeastern Caribbean, because nonreef assemblages contained virtually none of the epifaunal taxa that carried the heaviest damage in reef assemblages. An unambiguous test was not possible, however, because an unknown portion of the damage to epifauna might have occurred during life rather than after death (taphonomic signatures were based on damage to both interior and exterior surfaces of shells). Studies that have evaluated a single "target taxon," either to control for supply effects or as an exemplar of an entire class of shells, have yielded mixed results: some taxa show little environmental variation in damage (Dent 1995; Gardiner et al. 1995), other taxa are highly variable (Meldahl and Flessa 1990; Cutler 1995), and different taxa may vary in different ways across the same suite of environments (e.g., Fiege and Fürsich 1991; Dent 1995; Pandolfi and Greenstein 1997). In light of these mixed outcomes, the magnitude and nature of the effect that shell supply has on assemblage-level damage profiles deserves fuller testing.
Here, we test the effect of intrinsic biological factors on preservation in five tropical subtidal environments of the Bocas del Toro embayment in Caribbean Panama (Fig. 1). In our taphonomic survey of this area (Best and Kidwell this issue), we found that levels of damage were greater in bivalve death assemblages from environments with hard substrata than in exclusively soft-sediment environments, and that differences among environments within these sets were relatively subtle. A combination of extrinsic environmental differences probably drives this basically dichotomous pattern, including differences in rapidity and continuity of shell burial, and in the effectiveness of burial in protecting shells from various foulers and indeed in creating less tolerable conditions for foulers (linked to sediment grain size). Field experiments are being analyzed to test the roles of specific environmental conditions. However, the striking differences in death-assemblage condition between hard-substrate and soft-sediment environments may be in part a consequence of differences in shell supply, since hard-substrate assemblages contain a much larger proportion of epifaunal bivalves (average 42% versus 6% respectively; Table 3 in Best and Kidwell this issue). Epifaunal lifestyles among bivalves are stereotypically linked to calcitic shell compositions and thick shells, which in laboratory experiments are correlated with greater shell durability (e.g., Chave 1964; Flessa and Brown 1983; review by Kidwell and Bosence 1991).
Bocas death assemblages are especially advantageous for such a test:
Except for modest amounts of shell transported from patch reefs into immediately adjacent muds (≤∼10 m laterally), shell supply is derived from the local benthic community (Best and Kidwell this issue); thus, facies-level differences in damage are not smeared by large quantities of exotic shells.
Epifaunal bivalves are reasonably abundant in death assemblages from four of the environments and infaunal bivalves are abundant in all five (Best and Kidwell this issue). Death-assemblage compositions are thus not so strongly segregated to prevent cross-environment comparisons of the effect of life habit.
Intrinsic attributes of interest are not overwhelmingly intercorrelated in this bivalve fauna: (a) both infauna and epifauna comprise approximately equal numbers of thin-shelled and thick-shelled taxa, and, (b) although all shells composed of foliated calcite are from taxa with epifaunal life habits, an equal number of epifaunal taxa and the majority of epifaunal individuals have shells that are entirely or partly aragonitic (Table 1).
Only shells >8 mm were analyzed and the vast majority fell within the 8–20 mm range. Thus, any size effects are minimized.
Finally, in constructing damage profiles, only damage to the interior surfaces of shells was considered, and thus all encrustation, boring, commissural edge-rounding, and fine-scale alteration of shell material are unambiguously postmortem in origin.
Bocas del Toro is a large, geomorphically complex embayment of the Caribbean coast of Panama at 9 degrees North (Fig. 1; see Best and Kidwell this issue for more detailed description). Subtidal habitats are dominated by siliciclastic sands and muds shed from the Panama mainland and from islands within the embayment. Patch reefs rise up from these muddy seafloors and, together with algal meadows, constitute point sources of pure carbonate sediments. Death assemblages were collected from five sedimentary environments in November 1994 (same samples and stations as Best and Kidwell this issue; stations are nonconsecutively numbered polygons in Fig. 1). Three environments include some proportion of hard substrata: (1) patch reefs (primarily hard bottom of live and dead coral and coralline algae, with pockets or veneers of shelly carbonate sand; sampled at 14–16 m depth); (2) Halimeda-rich gravelly sand (coralgal meadows swept by strong currents and containing trace amounts of siliciclastic sand, sampled at 12–13 m depth); and (3) siliciclastic to mixed carbonate-siliciclastic mud among closely spaced patch reefs (bioclastic grain content >2 mm varies with proximity to reefs, sampled at 14–20 m depth). The other two environments are exclusively soft-sediment habitats (sampled at 13–29 m depth): (4) siliciclastic sandy mud (from wave- and current-affected margins of Canal del Tigre) and (5) homogeneous siliciclastic clayey mud (backwaters of Bahía Azul, and extensive submarine plain of innermost Laguna de Chiriquí, which has 37 m maximum depth).
Taphonomic data were collected from bivalve shells and fragments >8 mm. Damage profiles for the entire bivalve assemblage are based on the six taphonomic variables showing greatest variation. These are (1) disarticulation (presence versus absence); (2) fragmentation (versus unbroken); and, using shell interiors only and 10× magnification, (3) encrustation (presence versus absence; plus taxonomic composition of encrusters); (4) nonpredatory boring (presence versus absence; plus taxonomic composition of borers); (5) rounding of commissural edges (versus pristine, chipped, or thinned); and (6) fine-scale alteration of the shell's surface texture outside the pallial line (surface chalky and pitted, or eroded, versus pristine, dulled, chalky, or pitted; SEM required to determine specific cause, e.g., dissolution of crystallites versus maceration of organic matrix versus microboring).
For this study, each bivalve specimen was also identified taxonomically and scored for intrinsic attributes: bivalve life habit (epifaunal, infaunal, semi-infaunal; i.e., likely position at time of death), attachment (free-living, byssate, cemented, boring), shell thickness (> or < 0.5 mm), and shell mineralogy and microstructure (low-organic foliated calcite, low-organic porcellaneous aragonite, low-organic bimineralic, high-organic aragonite, and high-organic bimineralic). Information on bivalve shell compositions at the family level was derived from many sources, summarized in Kidwell and Brenchley 1996.
Each of the five sedimentary environments in the embayment yields a distinctive damage profile (taphonomic signature; Best and Kidwell this issue) (Fig. 2). Disarticulation and fragmentation are high across all environments except patch reefs, where many specimens were found still cemented to or wedged within reef rock (rubble retrieved by dredge). Because there is no correlation between damage levels and either water depth (range 12–29 m) or inferred water energy (relatively exposed stations 1, 7, 10, and 11, versus lee stations 5, 8, 12, and 13, in Fig. 1) Best and Kidwell (this issue) concluded that fragmentation and disarticulation were probably biogenic rather than physical in origin. This damage may, in fact, be largely due to predation rather than to strictly postmortem processes.
The other four variables in the damage profile (encrustation, boring, edge-rounding, and fine-scale alteration) are strictly postmortem in origin because they were measured from shell interiors. Damage of these types is significantly higher in hard-substrate environments than in exclusively soft-sediment environments. Damage profiles of environments differ either (1) in the specific ranking of variables (e.g., fine-scale alteration and encrustation are tied for top rank in the patch reef profile, but are ranked 1 and 2 in both Halimeda gravelly sand and in mud among patch reefs) or (2) in the magnitudes of similarly ranked variables (e.g., assemblages from homogeneous muds have significantly less encrustation, boring, and edge-rounding than those from sandy muds; 95% confidence interval [CI]). Preliminary scanning electron microscopy indicates that edge-rounding and fine-scale surface alteration are caused primarily by biological attack (microbioerosion, maceration), as of course are encrustation and boring.
Experimental work by many workers, both in the lab and field (reviewed by Kidwell and Bosence 1991; Parsons and Brett 1991), as well as our ongoing experiments in Panama (Best 1998; Best and Kidwell 1996), indicate that, although microbial attack continues during burial, other kinds of damage accrue either exclusively or most rapidly during shell exposure on the seafloor. Therefore, key extrinsic environmental factors differentiating taphofacies within the Bocas del Toro embayment were probably (1) the delayed burial or more frequent exhumation of shells in hard-substrate environments (i.e., greater total residence time on or just below the sediment-water interface in relatively coarse and permeable sediments) and (2) the rapid and more constant cover of shells by (siliciclastic) muds in the exclusively soft-sediment environments, which are also less favorable habitats for light-sensitive fouling taxa (Best and Kidwell this issue).
Effect of Life Habit.
Operationally, "epifauna" includes all taxa that live largely or entirely above the sediment-water interface. Such species cement themselves to hard substrata ("cementers"), are byssally attached to or nestle among hard substrata, live unattached upon the seafloor (free-living), bore into hard substrata, or live semi-infaunally. That is, "epifauna" includes all taxa that, upon death, would ordinarily lie on or largely above the granular sediment-water interface, accessible to overlying water and the array of taphonomic processes that operate there. "Infauna" refers only to bivalves that live fully within sediments, and thus, upon death, would ordinarily lie within the sedimentary column, requiring postmortem exhumation to be exposed to overlying water. Overall, sampled Bocas death assemblages comprise 37 epifaunal taxa and 49 infaunal taxa, totaling 407 and 1146 individuals respectively (Table 1).
Within individual samples, both cementing and noncementing epifaunal bivalves show higher than expected frequencies of encrustation and nonpredatory boring, given their abundance in the total death assemblage, and infauna consistently show lower than expected levels (Fig. 3).
Damage profiles of the five environments based on pooled samples are presented in Figure 4; all taphonomic variables are plotted except disarticulation, which is high in all environments. Both epifauna and infauna show the same environmental pattern that is evident in the total assemblage (Fig. 2): damage levels are significantly higher in hard-substrate environments (patch reef, Halimeda gravelly sand, mud among patch reefs) than in exclusively soft-sediment environments (sandy mud, mud). The most significant difference between epifaunal and infaunal shells is that, within hard-substrate environments, epifaunal shells show significantly higher frequencies of all kinds of damage (∼20 percentage points higher) than infaunal shells. (The only exception is the significantly lower fragmentation of epifaunal shells in patch reefs—many of which are still attached or wedged within the reef-rock framework—in contrast to infauna, which show high fragmentation in all environments.) Consequently, the taphonomic contrast between hard-substrate and exclusively soft-sediment environments is much greater when based on epifaunal shells than on infaunal shells (difference of ∼30 percentage points versus ∼10 percentage points).
Although epifauna show higher overall frequencies of damage than infauna in hard-substrate environments, the rank order of postmortem variables—that is, the pattern of damage—is identical or nearly so (Fig. 4). In Halimeda gravelly sand, the rank order of variables for epifaunal shells is surface degradation (most frequent damage), encrustation, edge rounding, and boring (SD > ENC > ER > BOR), and, within the narrower range of values for infauna, SD > ENC = ER > BOR. In mud among patch reefs, the epifaunal profile is SD > ENC > ER ≥ BOR, and for infauna, SD > ER ≥ ENC > BOR. In patch reefs, the epifaunal order is SD = ENC > ER = BOR, and for infauna, SD = ENC = ER = BOR because of large error bars. In sandy mud and mud, epifaunal shell damage is not significantly different from zero for any variable except SD in sandy mud; among the far more numerous infaunal shells, the only non-zero variables in either environment are SD = ER. Given the number of comparisons, this slight variation in rank orders is expected by chance.
Thus, with the exception of fragmentation in patch reefs, epifauna show significantly higher frequencies of all types of damage than infauna in the three hard-substrate environments, but the basic taphonomic profile (ranking of types of damage) is very similar to that of infauna.
Effect of Bivalve Shell Thickness.
With the exception of fragmentation, thick shells (>0.5 mm) show significantly higher levels of damage in hard-substrate environments (patch reef, Halimeda gravelly sand, mud among patch reef) than in soft-sediment environments (sandy mud, mud; frequencies of ∼30–70% versus <10%, Fig. 5A). This is the same broad trend seen in the total death assemblage (Fig. 2). Thin shells (<0.5 mm) exhibit the same trend (Fig. 5B), but differences are muted (20–40% damage frequencies in hard substrata versus ∼10% in soft-sediment) and not all pairwise comparisons are significant. For example, surface degradation is significantly higher for thin shells only in the Halimeda gravelly sand and in mud among patch reef environments, and edge-rounding is significantly higher only in mud among patch reefs. Both thick and thin shells show significantly lower fragmentation in patch reefs than in other environments.
Comparing thick and thin shells within each environment, the most notable difference is within hard-substrate environments, where thick shells show much higher damage frequencies than do thin shells (∼20 percentage points higher). In detail, thin shells are significantly less fragmented than thick shells in patch reefs, and significantly more fragmented in mud among patch reefs, sandy mud, and mud. The rank ordering of taphonomic variables in each environment is otherwise generally the same for both thick- and thin-shell data sets. In patch reef environments and Halimeda gravelly sand, CIs for thin shells are too large given the low frequencies to rank variables confidently, but strong similarities to the respective thick-shell rankings are evident: in patch reefs, surface degradation and encrustation are most frequent, followed by edge-rounding and boring (SD = ENC > ER = BOR), and in Halimeda gravelly sand, both thick and thin shells have profiles with SD > ENC > ER > BOR. In mud among patch reefs, thick- and thin-shell profiles differ in the order of only one variable, encrustation, which drops from second to third position: for thick shells, SD > ENC > ER = BOR, whereas for thin shells, SD > ER > ENC > BOR. In sandy mud, both thick- and thin-shell taxa show SD = ER > ENC = BOR, and in mud, both have profiles SD = ER > ENC = BOR = 0.
Thus, with the exception of peri-mortem fragmentation, thick shells exhibit significantly higher frequencies of damage than thin shells in the set of hard-substrate environments, but the basic pattern of damage is otherwise extremely similar to that of thin shells in both these and soft-sediment environments.
Effect of Shell Mineralogy and Microstructure.
To test the effect of shell composition on damage profiles, we grouped taxa into three categories (Fig. 6). Operationally, "calcitic bivalves" include all shells composed of foliated calcite, which is the relatively low-organic calcitic microstructure used by pectinids, ostreids, and, in combination with low-organic porcellaneous aragonite, by limids, spondylids, and plicatulids. Because foliated calcite forms the outer shell layer in these bimineralic taxa, and thus the area outside the pallial line on shell interiors, we group these bimineralic taxa with entirely calcitic taxa. This calcitic category constitutes 10% of all shells (Table 1). "Porcellaneous aragonitic bivalves" include calcite-free shells dominated by low-organic aragonitic microstructures such as cross-lamellar and complex cross-lamellar, and dominate all assemblages (80% of all shells; Table 1). This includes most infaunal bivalve taxa sampled in Bocas environments, and also many epifauna, such as chamids, arcids, anadarids, barbatiids, gastrochaenids, hiatellids, and lithophagids. The third category comprises all "high-organic shells," whose variants are unevenly distributed among environments. These include (1) entirely aragonitic pandorids composed of high-organic nacreous and homogeneous aragonites, (2) nuculanids, whose shells are interleaved mixtures of porcellaneous aragonite and conchiolin (the only high-organic taxon in sandy mud and mud environments), and (3) bimineralic taxa such as isognomids and pinnids, which combine nacre and high-organic prismatic calcite (exclusively in hard-substrate environments) (sources of information on microstructure types in Kidwell and Brenchley 1996).
The basic trends of the total death assemblage are captured by both the foliated calcite and the porcellaneous aragonite subsets of taxa: both subsets show higher frequencies of damage in hard-substrate environments than in soft-sediment environments (Fig. 6A,B). Again, the only exceptional variable is fragmentation, which for porcellaneous aragonite shells is high in all environments except patch reefs. CIs for the foliated calcite data are too large to detect any trend in fragmentation. Within each soft-sediment environment, calcite and porcellaneous aragonite shells show no significant differences in frequencies of damage (Fig. 6A,B): all types of damage other than fragmentation are low (≤ ∼10%).
Within each hard-substrate environment, pairwise comparisons of damage to calcitic and aragonitic shells mostly yield no significant or consistent differences. In patch reefs, encrustation and surface degradation are marginally lower for calcitic shells than for aragonitic shells, but in mud among patch reefs, encrustation and edge-rounding are marginally higher. The strongest divergence is in Halimeda gravelly sand, where encrustation, surface degradation and edge-rounding (but not boring) are significantly higher for calcitic shells than aragonitic shells. Despite these slight differences in frequency, the relative ranking of damage types for the two shell types does not differ significantly. In patch reefs, the ranking is ENC = SD > BOR = ER for both groups (with aragonite having marginally higher overall frequencies); in Halimeda gravelly sand, the ranking is SD > ENC > ER > BOR for both groups (with calcite having higher overall frequencies); and in mud among patch reefs, the ranking is SD > ER = ENC > BOR (with calcite having marginally higher overall frequencies).
Assessing the effect of organic content is less straightforward. (1) High-organic shells in sandy mud and mud (all pure-aragonite nuculanids) show no significant differences in frequency of damage compared with counterpart low-organic porcellaneous aragonitic shells in these environments (compare Fig. 6C with Fig. 6B), with the exception of significantly higher levels of edge-rounding and surface degradation among high-organic shells in sandy mud. (2) High-organic shells in patch reefs and mud among patch reefs (dominantly bimineralic forms) show lower frequencies of damage than counterpart low-organic foliated calcite shells (compare Fig. 6C with Fig. 6A), whereas there are no significant differences within Halimeda gravelly sand environments.
Thus, with the caveat of relatively large CIs, both low-organic foliated calcite and low-organic porcellaneous aragonite shells show greater damage in hard-substrate than in soft-sediment environments, consistent with the trend in the total bivalve death assemblage. Within environments, neither group shows a consistently higher frequency of damage than the other does, and among hard-substrate environments the two groups exhibit identical patterns of damage (ranking of damage types). High-organic microstructures show less damage than low-organic microstructures of the same mineralogy in two of three hard-substrate environments, but greater damage than low-organic counterparts in one of the two soft-sediment environments. Thus, mineralogy and microstructure convey no consistent effect on taphonomic damage.
Damage to shell interiors (encrustation, boring, edge-rounding, and fine-scale surface degradation) is unambiguously postmortem in origin. Patterns of damage within Bocas bivalve assemblages thus permit us to determine the roles of extrinsic and intrinsic taphonomic factors in generating facies-level differences in shell damage, and to discriminate links between postmortem damage and specific intrinsic characters (life habit, shell thickness, shell mineralogy and microstructure). As a complement to experiments that measure the reactive rates of shell types, our results indicate the net product of differential shell reactivities in a time-averaged system.
Our key findings are: (1) each subset of the bivalve fauna (four pairwise divisions: epifaunal versus infaunal life habit, thick versus thin shell, foliated calcite versus porcellaneous aragonite, and high- versus low-organic microstructures) displays greater damage in hard-substratum environments than in soft-sediment environments, as also observed in the pooled data set (Fig. 2); (2) in any given environment, all subsets exhibit the same or a nearly identical pattern of damage in terms of the rank order of taphonomic variables; and (3) subsets of shells differ in damage from one another, if at all, only in the percentage of shells affected (Table 2).
These empirical results have a series of implications, both for the behavior of taphonomic systems and for the sedimentologic and paleobiologic value of death assemblages.
Relative Importance of Extrinsic and Intrinsic Factors.
In Bocas del Toro, extrinsic environmental conditions exert an overwhelming effect on death assemblage condition: all [biological] subsets of the bivalve death assemblage of a given environment exhibit the same basic pattern of damage (Table 2). Intrinsic effects were detected only in the three environments containing hard substrata, where the composition of shell supply modulates the strength of the taphonomic signature: epifaunal shells and thick shells exhibit the same basic patterns of postmortem damage as their infaunal and thin-shelled counterparts, but do so with significantly higher frequency. The larger the proportion of epifaunal and thick shells, the more strongly the taphonomic signature of the environment is registered in the total death assemblage.
Relative Importance of Different Intrinsic Factors.
The intrinsic factors that are most strongly linked to high frequencies of shell damage are epifaunal life habit and shell thickness (>0.5 mm), which covary in Bocas death assemblages: 72% of epifaunal shells are also thick, whereas only 41% of infaunal shells are thick (Table 1). Mineralogy does not appear to be a factor: only 36% of epifaunal shells are composed of foliated calcite (an additional 11% are composed of prismatic calcite; data from Table 1). In general, shell damage within Bocas bivalve assemblages is not linked to shell mineralogy: among low-organic microstructures, foliated calcite conveys a greater likelihood of shell damage only in the relatively high-energy Halimeda gravelly sand environment (Fig. 6A,B).
Our results suggest that high-organic microstructures may be more of a preservational liability among infaunal taxa (or in fine-grained substrata) than among epifaunal taxa (or in coarse and hard substrata), but these microstructural types are uncommon in all environments and much larger sample sizes are needed to test the effect with confidence. In future studies, the additional sampling effort would be well worthwhile: although high-organic microstructures are relatively uncommon in Recent molluscan faunas, they constitute an increasing proportion of bivalve faunas back into the fossil record.
Causes of Intrinsic Effects.
We attribute the higher frequency of damage among epifaunal shells to their death above the sediment-water interface, which creates an opportunity for postmortem modification immediately after death, especially by encrustation and boring, that typically does not exist for infaunal shells. It is interesting that, even in hard-substrate settings where all shells probably undergo multiple cycles of burial and exhumation on the seafloor, and where some epifaunal individuals may even be buried alive, epifaunal shells are still damaged more frequently than infaunal shells. This suggests that taphonomic processes are operating rapidly, such that even a short initial delay in burial can launch epifauna significantly ahead of infauna in the accumulation of taphonomic damage. Our ongoing experiments in Caribbean Panama suggest in fact that significant damage occurs within months of exposure in all environments (Best and Kidwell 1996; Best 1998).
The higher frequency of damage to thick shells may reflect the preferential selection by infesting taxa of the most durable (bulky) shell substrata in a local death assemblage, regardless of whether these shells are exposed at the sediment-water interface immediately after death or at some later time. Highly selective settlement behavior by encrusting and boring organisms is well documented in both the ecological and paleontological literature, including selective infestation of the thickest portions of live and dead shells. This kind of behavior might also contribute to the high damage frequencies observed among epifaunal shells if, for reasons of shell microstructure, infesters preferentially colonize the interiors of epifaunal shells (e.g., microbial preference of nacre over porcellaneous aragonite observed by Glover and Kidwell [1993]).
The higher frequencies of damage among epifaunal and thick shells, and the lack of consistent taphonomic differences between calcitic and aragonitic shells, may seem to contradict timed-release experiments (e.g., shells tumbled with abrasive, submerged in acid, exposed to infesters [Chave 1964; Driscoll 1970; Peterson 1976; Flessa and Brown 1983; Glover and Kidwell 1993]). Such studies consistently find that different shell types accrue damage and disintegrate at different rates, and that the classes of bivalve shells having lowest reactivities are calcitic (all of which are epifaunal species), thick (small surface area to mass ratio), low in intraskeletal porosity, low in microstructural organic content, and large bodied. However, in time-averaged assemblages, where shells have been supplied throughout an interval of time rather than at only a single moment, the rapid destruction of weak shells should result in (1) "durable" subsets of taxa exhibiting more rather than less damage than fragile subsets (basic pattern observed in Bocas) and (2) durable subsets having greater degrees of time-averaging (i.e., the widest range in shell age-since-death). This latter point has not yet been tested in Bocas via direct-dating. Elsewhere, direct-dating of Holocene bivalves has shown that, although damage accrues episodically rather than continuously, older shells within time-averaged assemblages are more consistently in poor condition than younger shells (Meldahl et al. 1997).
Thus, in addition to the probable effects of (1) an initial window of postmortem exposure on the seafloor not shared by all shell types and (2) preferential attack on some shell types by taphonomic agents, the observed high damage to epifaunal and thick shells might also reflect (3) some degree of disharmonious time-averaging (sensu Kowalewski 1996), with some biological subsets of the death assemblage (durable epifaunal and thick-shelled taxa) having greater average shell age-since-death than others.
Comparison with Previous Studies.
Parsons and Brett (1991) also noted that epifaunal shells had higher levels of damage than infaunal shells in reef sediments, but because their data combined damage to both shell exteriors and interiors, the taphonomic contribution to the pattern was unclear (the exterior surfaces of epifaunal shells may be damaged during life). Within a series of pure-carbonate subtidal environments of southern Florida, Dent (1995) also reported higher levels of encrustation and boring for two epifaunal target taxa (chamid bivalves, cerithiid gastropods) than for an infaunal taxon (tellinid bivalves), whose damage was limited to edge alteration, surface degradation, and abrasion (damage to all surfaces pooled).
Studies that target damage to single taxa aim either to control the variable of shell supply on taphofacies or to use that taxon as an exemplar of a class of shells, and thus comparisons to our results are not straightforward. These studies also pool damage to shell exteriors with data on shell interiors, and thus are testing the overall condition of shells rather than strictly postmortem damage. Most studies find that target taxa collected from different environments have different taphonomic signatures, demonstrating that extrinsic factors can be significant (Meldahl and Flessa 1990; Fiege and Fürsich 1991; Cutler 1995; Pandolfi and Greenstein 1997).
In a single environment, however, these studies show that different target taxa can yield very different frequencies and profiles of damage, suggesting the potential for significant intrinsic supply effects. Dent (1995), for example, found that the damage profiles of one epifaunal target taxon were invariant across south Florida environments (chamid bivalves; probably allochthonous in some environments), a second epifaunal taxon (gastropod Cerithium) closely resembled the total death assemblage in each environment, and an infaunal target taxon (tellinid bivalves) varied in damage among environments but diverged from the total assemblage. Our analysis of complete subsets of taxa reveals the magnitude of these potential supply-effects and finds that the intrinsic nature of local bivalve communities has only quantitative rather than qualitative effects on overall death assemblage condition in tropical subtidal environments.
Our study does not contradict earlier work, but is the next logical step, and it underscores the importance of testing for relative effects of intrinsic and extrinsic factors, to determine if, for example, intrinsic factors are more important in some depositional systems than in others. Biological subsets of assemblages, defined by life habit or by construction (e.g., pelmetazoan study by Meyer et al. 1989), are probably more meaningful units of comparison than single taxa for such analyses, and would maximize the confidence of extrapolating results into the fossil record. This approach could be used in any setting but is especially practical in tropical systems where species richness and equitability are typically high, making it difficult to acquire statistically meaningful numbers of single target species in all environments.
Analysis of bivalves from Caribbean Panama indicates that in each of the environments studied, all biological subsets exhibit the same pattern of damage (rank order of damage types) and differ only in the absolute intensity of that damage. This result has several advantages for taphofacies analysis. First, it means that shell supply has a quantitative rather than a qualitative effect on taphofacies—the nature of the local bivalve community modulates the intensity of the damage style, not its fundamental pattern. In this situation, a residual assemblage of calcitic taxa or molds of thick shells would yield the same basic environmental signature as the complete bivalve fauna, were it preserved. Second, some subsets of the fauna, particularly epifauna or thick shells, could be targeted for facies analysis because they register taphonomic differences more strongly than other death-assemblage components.
From a paleobiologic perspective, it is not clear whether these within-assemblage differences in the intensity of damage (seen only in hard-substrate environments) signify differences in degree of time-averaging (range in age-since-death of individuals), exposure time (which may be ≤ age-since-death), selectivity in attack by taphonomic agents, or some combination of these factors, which have different consequences for bias in paleobiologic information (loss in time resolution, versus modification of species' relative abundances). These issues are under investigation using a combination of field experiments and direct-dating (Best unpublished; Best and Kidwell unpublished; Best et al. unpublished). The implication is clear, however, that these subsets of the faunal record, although from the same environment, are probably not iso-taphonomic in terms of bias.
The next question is the generality of our results. Existing data sets on the condition of death assemblages in other study areas could be dissected into biological categories like those here, and thus might rapidly yield comparable information on the effect of shell supply in other settings. A priori, it is difficult to anticipate how—or whether—patterns of damage accrual in death assemblages vary geographically without some form of empirical data. For example, extrinsic environmental effects may dominate over much of the tropical shallow-marine seafloor because, as in Bocas, biological agents play an extremely large and aggressive role in shell destruction (encrusters, borers; also elevated rates of microbial maceration? Best and Kidwell this issue). On the other hand, within the Bocas study area, it is hard-substrate environments that host death assemblages with the greatest damage. Although this reflects a combination of extrinsic and intrinsic factors (frequent reworking, low net burial rates, abundant epifauna), some of these same conditions would apply on cobble-, rock-, and shell-grounds in higher latitudes, and in fact molluscan assemblages there can comprise highly damaged material (e.g., Wilson 1982; Young and Nelson 1988). That is, among shallow subtidal environments, perhaps grain size or low net sedimentation, and the other biological and physical conditions linked with it, are the key variables rather than depositional system per se (carbonate versus siliciclastic) or latitude and climate.
A systematic exploration of major environmental settings, whose assemblages are dissected into intrinsic biological categories such as life habit and shell construction for analysis, is thus required to determine the overarching patterns of taphonomic damage. Accompanied by field and laboratory tests of causality, including how damage and bias correlate, we will then be better positioned to extrapolate back into the fossil record, where the proportions of benthic life habits and hard-part types have changed over evolutionary time.
We thank A. I. Miller, G. H. Cadée, and D. Jablonski for helpful reviews of the original manuscript. Supported by National Science Foundation grant EAR-9628345.
- Accepted 11 June 1999.


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Shell-operated production rates that remain shut in as a result of Hurricane Dean are approximately 30,000 barrels of oil per day and 70 million cubic feet of natural gas per day. We are beginning to bring production back on line.
We continue to monitor Dean closely, as well as other activity in the tropics.
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