Assessing Benthic Feeding Strategies in Continental Margin

Environments Using Radiocarbon

David J. DeMaster*, Craig R. Smith+, & Carrie J. Thomas*

*
Dept. of Marine, Earth and Atmospheric Sciences, North Carolina State University, Raleigh,
North Carolina 27695, USA
+
Dept. of Oceanography, SOEST, University of Hawaii, 1000 Pope Road, Honolulu, Hawaii
96822, USA

Keywords: Biogeochemistry, radiocarbon, benthic ecology, marine carbon cycling

Surface-deposit feeding megafauna, such as holothurians, play an important role in

continental margin ecosystems because they ingest and assimilate much of the labile

organic matter deposited on the seafloor prior to extensive microbial processing or

ingestion by coexisting infauna. Radiocarbon (or carbon-14) measurements are a useful

tool in determining if labile organic material is reaching the seabed and in resolving feeding

strategies for deposit-feeding benthic fauna living on the continental margin because

enrichments in this isotope (produced primarily by stratospheric nuclear bomb testing)

signify freshly formed, labile organic carbon. In Santa Catalina Basin off California and

on the Antarctic continental margin carbon-14 data indicate that labile material is reaching

the seabed and that deposit feeders are very selective as they ingest and assimilate this

organic matter. Here we show that some megafaunal deposit feeders such as the surface-

deposit-feeding holothurian, Peniagone sp., primarily attain their carbon-14 enrichment

(i.e., their labile organic matter) via particle selection during the ingestion process, whereas

1
other benthic fauna such as the subsurface-deposit-feeding holothurian, Molpadia

musculus, primarily attain their enrichment via selective digestion and other assimilative

processes.

Lauerman et al.1 used sediment ingestion rates to estimate that as much as 4% of the vertical

mass flux reaching the seabed in the abyssal northeast Pacific Ocean is ingested by a single

epifaunal holothurian species (Abyssocucumis abyssorum). In Santa Catalina Basin (water depth

1240m) Miller et al.2 developed a 234Th flux model and determined that the 3 dominant

megafaunal surface deposit feeders consume 40-50% of the daily flux of organic material

reaching the seabed. These studies clearly indicate that large, seemingly sparse surface-deposit

feeders can have a substantial impact on the fate of organic matter reaching the seabed, and

consequently, on food-web dynamics and sediment biogeochemistry.1,2,3,4,5 One disadvantage of

using sediment processing rates and particle-reactive tracers (such as 234Th) to study deposit-

feeding processes is that the bulk sediment weights and tracers do not distinguish the dominant

sources of organic matter sustaining benthic ecosystems. Here we highlight the use of naturally

occurring 14C as a tracer of labile organic matter as it moves through benthic food webs. In

addition to elucidating the nature of organic matter incorporated by deposit feeders, this naturally

occurring tracer provides important insight into the feeding strategies used to obtain labile

organic carbon in habitats dominated by refractory organic material.

During the late 1950’s and early 1960’s, nuclear-bomb testing in the stratosphere

enhanced the production of radiocarbon in the atmosphere. This enrichment has been detectable

in surface marine plankton during the past 4 decades, providing a very useful tracer of recently

produced, labile organic matter. ∆14Corg valuesa for plankton collected near the ocean surface

2
changed from –90 - ~0 per mil prior to nuclear testing to +50 - +195 per mil6 after testing,

except in areas such as high latitudes where upwelling exceeds air-sea gas exchange. The 14Corg

contents at the surface of continental-margin deposits are affected by this planktonic signal as

well as by the 14Corg content of other sources of organic matter reaching the seabed, including

older, reworked marine and terrestrial organic material7. Benthic deposit feeders preferentially

incorporate the more recently produced labile organic matter into their tissues and the

radiocarbon measurements provide a tool for tracking these selection and enrichment processes.

The pioneering efforts using 14Corg to study continental-margin and abyssal food webs

yielded conflicting results. Pearcy and Stuiver8 measured radiocarbon in benthic fish and large

invertebrates from the Oregon continental margin (~ 2800 m water depth) and concluded that

recently produced (rapidly sinking) particles from the surface ocean were not the major source of

nutrition to deep-sea benthos. In contrast, Williams et al.9 concluded, based on 14C

measurements, that the main source of dietary carbon for meso-, bathy-, and abysso-pelagic fish

and crustacea was rapidly sinking organic detritus from the surface ocean. More recently,

DeMaster et al.10 used radiocarbon analyses to determine whether recently produced, labile

marine organic carbon was deposited on the seafloor at 800m water depth on the North Carolina

continental slope. The ∆14C value of the bulk organic matter in near-bottom particle traps from

the Mid-Atlantic Bight ranged from –120 to +4 per mil11,12, suggesting that little or no bomb-

produced 14C was reaching the seabed. However, benthic fauna from the North Carolina margin

had ∆14Corg values as high as +82 per mil and averaged +56 per mil (SD = 20; n=10); i.e.,

________________________
a 14
∆ Corg (in per mil) = [Activitysample/(0.95 *Activitystandard) – 1] * 1000, where the Standard
(NBS Oxalic Acid) and Sample activities (in dpm/gm C) are corrected for isotopic variations
based on 13C content. The “org” subscript denotes that the form of carbon is organic matter.

3
substantially higher than the ∆14C contents of organic matter in surface sediments (–215 to –41

per mil)10. Based on mass-balance calculations, the percentage of freshly produced, labile

marine organic carbon in the upper few centimeters of the seabed was only ~0.2% of the total

organic carbon (or approximately 0.0034 wt. % of the total sediment), yet this labile carbon was

the predominant source of nutrition for the deposit-feeding macrobenthos (as evidenced by the

high bomb-14C contents in the animal tissues). 14

C enrichment in deposit-feeder tissue is likely

to occur via two processes13,14: (1) selection of particles rich in labile organic material during the

process of ingestion (i.e., “selective feeding”); and (2) preferential digestion and/or assimilation

of recently-produced, labile organic matter once it has entered the gut (i.e., “selective

assimilation”).

Study sites and methods

Our field sites were Santa Catalina Basin (California Borderland) and the western Antarctic

Peninsula continental shelf. Both sites have a variety of deposit-feeding megafauna, whose large

body sizes enable sampling of the necessary gut sediments and tissues. The Santa Catalina Basin

site (33º 9.5’N, 118º 28’W, 1240 m water depth) is off the California coast, where coastal

upwelling sustains a relatively high flux of organic carbon through the water column year

round15. The depositional environment is low energy because of the basin topography. Settling

material was collected 170m above the seafloor in a 0.16m2 conical particle trap, which was

sampled during each of 5 cruises between January of 1996 and April of 1998 (typically 60-70

day collection period). An 8m otter trawl was used to collect benthos from the seabed, and a

multiple corer was used to collect relatively undisturbed surface sediments. One sample of

4
surface-sediment floc was obtained during the April 1998 cruise by gently agitating a recently

recovered multiple core and then collecting the resulting suspended sediment.

Sampling on the west Antarctic Peninsula shelf was part of a program called

FOODBANCS (Food for Benthos on the Antarctic Continental Shelf), conducted approximately

100 km west of Palmer Station (65ºS, 64ºW). Three sites were established on the shelf in water

depths between 500 and 600 meters: an inner-shelf station (Sta. A); a mid-shelf station (Sta. B);

and an outer-shelf station (Sta. C). Two 0.16m2 conical particles traps, located 150m and 170m

above the seafloor, were placed on a mooring at Sta. B, near the site of the Palmer Station LTER

mooring16. An 8m otter trawl was used to collect benthos and a megacorer was used to collect

surface sediments during the 5 cruises, which occurred between November 1999 and March

2001. The sediments in both field areas (Santa Catalina Basin and the Antarctic Peninsula) were

dominated by mud-sized material, with organic carbon contents typically ranging from 3-4 wt. %

in Santa Catalina Basin and ~1% on the Antarctic shelf.

Gut contents and muscle tissue were dissected from individual megafauna (typically 5-15

cm in length) within hours of collection. Gut linings were excluded from gut-content samples

during dissections, except for echiuran worms, where the lining was included because of the
14
small diameter of the intestinal track. C activities were measured on the organic carbon

contents of the gut, which predominantly originated from ingested sediment, gut flora, and the

animal’s digestive fluids (the relative amounts of which were unknown). There is a potential for

lysis in gut cells as a result of pressure changes encountered during animal recovery; however,

animals undamaged by our trawling were alive upon recovery indicating that cell lysis is much

less likely to occur in our studies (water depths of 500-1200m) than in studies from abyssal

depths3. Gut and tissue samples were analyzed from individual animals except for the

5
holothurian, Peniagone sp., where samples were pooled from 4-5 individuals. Samples were

frozen on board ship, dried at 60ºC in the laboratory, and then leached with 1N HCl to remove
14
inorganic carbon. C abundance in the organic matter was measured on the Woods Hole

Oceanographic Institution accelerator mass spectrometer after combusting the organic matter on

a Carlo Erba 1108 CNS analyzer and cryogenically collecting the resulting carbon dioxide.

Stable carbon and nitrogen isotopic abundances were measured on organic samples using a C/N

analyzer-mass spectrometer system.

Results from Santa Catalina Basin (SCB)

Surface deposit feeders from SCB analyzed for tissue and gut 14C content included the epibenthic

holothurians, Pannychia moseleyi and Chiridota cf. pacifica, as well as the gastropod,

Bathybembix bairdii. The ∆14C activities from animal tissues (Figure 1) ranged from –2.6 to

+66.3 per mil, considerably enriched relative to the surface sediment floc activity (∆14C = -139

per mil) and the particle-trap samples (∆14C = -24 to –57 per mil). The particle-trap material

clearly has an old 14C component, probably derived from resuspended organic carbon from the

continental margin or from old terrestrial soil carbon6. The highest ∆14C activities in the animal

tissues were comparable to nearby living kelp 14C activity (+65.1 per mil), which was similar to

surface plankton values from southern California4. The 14C enrichments of megafaunal body

tissues from the California Borderland relative to the surface sediment (a ∆14C difference ranging

from +135 to +200 per mil) indicate that selective feeding and/or assimilative processes are

occurring not only on the North Carolina slope but in west coast marginal basins as well.

To investigate the processes responsible for 14C enrichments in deposit-feeder tissues, the
14
C activities in surface sediments, gut contents, and body tissues were compared in two

6
Pannychia moseleyi individuals collected in a single otter trawl. We considered differences in
14
C activities between surface sediment and the foregut material to result from particle selection

during deposit feeding, and the differences between 14C activities of foregut contents and body

tissues to indicate selective processes during digestion and assimilation. Pannychia moseleyi

individual #1 exhibited a 100 per mil 14C enrichment between surface sediments and foregut

contents (“particle selection”) and a 34 per mil enrichment between foregut contents and body

tissue (“digestive selection”, Figure 1). This individual clearly had ingested recently deposited,

labile organic matter as indicated by the high and down-gut decreasing organic carbon contents

(foregut, midgut, hindgut = 12.9, 8.7, and 7.7 weight %, respectively) and gut 14C activities

(foregut, midgut, hindgut = -37, -65 and –63 per mil, respectively). The trend in 14C activity is

consistent with the preferential digestion/assimilation of 14C-enriched labile organic matter in the

midgut. In P. moseleyi individual #2, the gut contents (∆14C = -119 to –144 per mil) were more

similar to the surface sediment floc (-139 per mil) indicating that this animal primarily had

ingested bulk surface sediment containing little or no fresh labile organic matter. The organic

carbon contents in the gut of P. moseleyi ind. #2 were relatively low and nearly constant

(foregut, midgut, hindgut = 5.6, 5.4, and 5.2 wt. %, respectively), suggesting little absorption of

organic material during gut passage. The lack of a trend in the down-gut 14C activities in

Pannychia moseleyi ind. #2 (foregut, midgut, hindgut = -136, -119, -144, respectively) is likely

to result from non-steady-state particle selection during feeding. For these holothurians the gut

throughput time is approximately 1 day2, whereas the body tissue integrates carbon assimilated

over a period of months to years, based on known growth rates of deep-sea echinoderms17.

Distributions of labile protein abundance can be patchy on meter scales in surface sediments of

Santa Catalina Basin due to heterogeneous bioturbation and depositional processes2,18.

7
Consequently, holothurians collected from a single otter trawl (which samples over 3-4 km) are

likely to have gut contents of different food value and lability (as has been observed in tropical

holothurians19). The variability in the 14C contents of the body tissues from the five Pannychia

moseleyi analyzed here (-3 to +66 per mil), however, suggest that even over time scales of

months to years (i.e., over the time scales of tissue growth and turnover) foraging success may

vary significantly among individuals within a single species of mobile, surface-deposit feeder.

We predict that growth rates and reproductive success also will exhibit substantial intraspecific

variability in surface deposit feeders studied at a single site. These findings demonstrate the

need to analyze multiple individuals from a particular species for their tissue and gut 14C

activities before a final conclusion is reached concerning feeding strategies and the relative

importance of particle selection and digestive selection processes.

Results from the Antarctic Continental Shelf

To determine whether particle- and/or assimilative-selective deposit feeding strategies are

common at high latitudes, where there is a dramatic seasonal change in food supply to the

benthos, 14C activities were measured in six species of megafaunal deposit feeders from the west

Antarctic Peninsula continental shelf (Figure 2). The surface-deposit feeders chosen for study

included the epibenthic holothurians, Peniagone sp., Protelpidia murrayi, and Bathyplotes sp.; a

head-down (conveyor-belt feeding) molpadiid holothurian, Molpadia musculus; an unidentified

echiuran worm; and a deposit-feeding urchin. The 14C contents of surface plankton for this area

typically range from –160 to –115 per mil20 because of the intense upwelling of older,

subsurface waters. The 14C contents of our particle trap material (∆14C = -165 to -141 per mil;

n=5) and most plankton tow material (-147 to -129 per mil; n=6, Figure 2) were consistent with

8
the regional surface plankton values from the literature20. One exception was the plankton

collected in March 2000 (∆14C = -36 per mil), whose high 14C abundance is believed to result

from an unusually intense air-sea gas exchange period. Although the 14C activities of the

Antarctic samples were significantly more negative than the California Borderland samples, the

trends among the ∆14C contents of surface sediments, gut contents and body tissues from

Antarctica were very similar to those from the California Borderland indicating the occurrence of

particle and digestive/assimilative selectivity in Antarctic megafaunal deposit feeders.

The 14C contents of the animal tissues were surprisingly uniform (∆14C = -136 to –113

per mil, n=10) considering the distances between sites (~100 km) and seasonal differences in

sampling (summer/winter). The animal tissues were considerably enriched in 14C (by 50 to 75

per mil) relative to the 14C content of the surface phytodetritus sample (-188 per mil), collected

during the summer of 2001. Presuming that the relative extent of particle selection can be

characterized by the ∆14C difference between surface sediment organic carbon and foregut

contents, there appears to be a systematic increase in particle selectivity in the summer data

comparing the molpadiid, urchin, Protelpidia murrayi, and Bathyplotes sp. to the Peniagone sp.

and echiuran worm (Figure 2). Molpadia musculus is the least selective benthos, with the 14C

difference between the surface sediment and foregut contents of only 8 per mil. Mopadiids are

head-down, conveyor-belt deposit feeders, and thus have reduced access to (14C enriched) labile

organic material at the sediment-water interface. The holothurian, Peniagone sp., and the

echiuran worm appear to be the most particle selective with their whole-gut contents enriched by

60 per mil in 14C relative to the surface sediment. Some of the 14C enrichment in the echiuran

worm gut may have resulted from the inclusion of the gut lining with the gut sediment sample.

Because the 14C activities of body tissues were relatively uniform across all six Antarctic species,

9
those animals characterized by the lowest particle selectivity (i.e., M. musculus, the irregular

urchin, Protelpidia murrayi, and Bathyplotes sp.) exhibited the largest digestive and/or

assimilative selectivity. These relatively non-particle selective deposit feeders exhibited

differences in 14C content between foregut and body tissue ranging from 42 to 49 per mil,

whereas the more particle selective Peniagone sp. and echiuran worm exhibited differences of

only 5 to 8 per mil (i.e., minimal digestive selection). The trends in particle selectivity and

digestive/assimilative selectivity also were evident in the winter data (Figure 2, top panel), which

showed that the enrichments between the foregut content and the body tissue of the molpadiid

(93 per mil), Protelpidia murrayi (78 per mil), and Bathyplotes sp. (67 per mil) were much

greater than in Peniagone sp. (essentially 0 per mil). In addition, the sequence of particle

selection efficiency for these taxa based on the 14C data matched that determined from the 234Th

content of the animal’s gut material21, where the number of analyses was much greater (n=48,

Figure 3) and seasonal coverage was more extensive.

When the 14C activities of gut contents in relatively non-selective particle feeders (i.e.,

molpadiid, Protelpidia murrayi, and Bathyplotes sp.) were compared seasonally (Figure 2), the
14
C content of the ingested organic matter generally was depleted during the winter relative to

the summer (typically by 40 per mil). This depletion is consistent with a reduced supply of labile

organic carbon (i.e., plankton detritus) reaching the seabed during the austral winter when

primary production becomes light limited and much of the ocean surface is covered with sea ice.

The highly particle selective Peniagone sp., however, was able to ingest relatively labile organic

matter (∆14C = -133 per mil) all year round, suggesting that a highly efficient foraging strategy

may have offset much of the seasonal variability in food quality and supply.

10
 The chemical differences among surface sediment, gut contents, and animal tissue were

apparent in other isotopic and elemental signatures besides 14C. Stable isotopes of carbon and

nitrogen and C/N weight ratios, all have been used to assess organic carbon sources and trophic

relationships within food webs22,23,24. In our study area, for example, the molpadiid tissue had a

δ13C, δ 15N, and C/N ratio of –20.5 per mil, +10.8 per mil, and 4.2, respectively, whereas the

molpadiid gut contents had δ 13C, δ 15N, and C/N values of –25 to-26 per mil, +4 to +5.2 per mil,

and 7.5, respectively. The dynamic range in these isotopic and elemental variations is

considerably smaller than with 14C, and the inference regarding lability is more direct with

radiocarbon than with the other tracers. These chemical data indicate that many deposit feeders,

including surface feeding holothurians, subsurface feeding holothurians, and irregular urchins

violate the isotopic adage “you are what you eat”. For those deposit feeders exhibiting strong

digestive/assimilative selection processes, it is more accurate to say: “You are what you

assimilate”. This latter formulation is likely to be appropriate for many types of detritivores

feeding on material of low food value25,26. Our 14C data indicate that deposit-feeding megafauna

are selectively incorporating labile organic matter into their tissues as a result of particle

selectivity during ingestion and/or preferential digestion/assimilation. Both strategies can

remove much of the energy-rich, labile organic material reaching the seabed prior to extensive

microbial decomposition in the sediment column.

11
References

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evidence for rapid ingestion of settling particles by mobile epibenthic megafauna in the abyssal
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6. Williams, P.M., Robertson, K.J., Soutar, A., Griffin, S.M., & Druffel, E.R.M. Isotopic
signatures (14C, 13C, 15N) as tracers of sources and cycling of soluble and particulate organic
matter in the Santa Monica Basin, California. Prog. Oceanogr. 30, 253-290 (1992).

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and particulate organic radiocarbon in the northwest Atlantic continental margin. Glob. Biochem.
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Sea Res. 30, 427-440 (1983).

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10. DeMaster, D.J., et al. Deposition of bomb 14C in continental slope sediments of the Mid-
Atlantic Bight: assessing organic matter sources and burial rates. Deep-Sea Res. II 49, 4667-4685
(2002).

11. Anderson, R., Rowe, G., Kemp, P., Trumbore, S. & Biscaye, P. Carbon budget for the mid-
slope depo-center of the Middle Atlantic Bight. Deep-Sea Res. II 41, 669-703 (1994).

12. Santschi, P.H., Guo, L., Walsh, I.D., Quigley, M.S., & Baskaran, M. Boundary exchange
and scavenging of radionuclides in continental margin waters of the Middle Atlantic Bight:
implications for organic carbon fluxes. Cont. Shelf Res. 19, 609-636 (1999).

12
13. Roberts, D. Deposit-feeding mechanisms and resource partitioning in tropical holothurians.
J. Exp. Mar. Biol. Ecol. 37, 43-56 (1979).

14. Lopez, G.R. & Levinton, J.S. Ecology of deposit-feeding animals in marine sediments.
Quat. Rev. of Biol. 62, 235-260 (1987).

15. Thunell, R.C., Pilskaln, C.H., Tappa, E. & Sautter, L.R. Temporal variability in sediment
fluxes in the San Pedro Basin, Southern California Bight. Cont. Shelf Res. 14, 335-352 (1994).

16. Smith, R.C., et al. The Palmer LTER: A long-term ecological research program at Palmer
Station, Antarctica. Oceanogr. 8, 77-86 (1995).

17. Gage, J.D. & Tyler, P.A. Deep-sea Biology: A Natural History of Organisms at the Deep-
sea Floor. Cambridge University Press, Cambridge (1991).

18. Demopoulos, A.W.J., Smith, C.R., DeMaster, D.J., & Fornes, W.L. Evaluation of excess
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Th activity in sediments as an indicator of food quality for deep-sea deposit feeders. Jour.
Mar. Res. 61, 267-284 (2003).

19. Uthicke, S. & Karez, R. Sediment patch selectivity in tropical sea cucumbers
(Holothurioidea : Aspidochirotida) analyzed with multiple choice experiments. J. Exp. Mar. Biol.
Ecol. 236, 69-87 (1999).

20. Gordon, J.E. & Harkness, D.D. Magnitude and geographic variation of the radiocarbon
content in Antarctic marine life – Implications for reservoir corrections in radiocarbon dating.
Quat. Sci. Rev. 11, 697-708 (1992).

21. McClintic, M.A. The delivery and fate of particles settling on the seabed adjacent to the
western Antarctic Peninsula: evidence from excess 234Th measurements. M.S. Thesis, North
Carolina State University. Raleigh, NC, 56 pp. (2002).

22. Deniro, M.J. & Epstein, S. Influence of diet on distribution of carbon isotopes in animals.
Geochim. Cosmochim. Acta 42, 495-506 (1978).

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18, 293-320 (1987).

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deposit-feeding and carnivorous polychaetes. Oecologia 82,1-11 (1990).

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26. Jumars, P.A., Mayer, L.M., Deming, J.W., Baross, J.A., & Wheatcroft, R.A. Deep-sea
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ACKNOWLEDGEMENTS: This research was supported by the National Science Foundation through the Chemical

Oceanography, Biological Oceanography, and Polar Ocean programs. Grant #’s:OCE-9527382 and OPP-9816049 to

C.R. Smith and D.J. DeMaster).

No Competing Interests.

Correspondence and requests for materials should be addressed to D.J.D. (e-mail: dave_demaster@ncsu.edu).

14
Figure Legends
14
Figure 1 C data from Santa Catalina Basin samples. The data show a clear enrichment in

organic carbon 14C activity between the surface sediment floc and the various animal tissues.

This enrichment occurs as a result of particle selection during ingestion as well as by preferential

selection of labile organic material during assimilation. The enrichment caused by particle

selection is designated as the activity difference between the surface sediment floc and the

animal’s foregut contents, whereas the assimilative selection is designated as the activity

difference between the foregut contents and the animal’s tissue. The two holothurians

(Pannychia moseleyi Ind. 1 and 2) were collected in the same otter trawl. Their gut contents

reflect a combination of labile carbon (as 14C rich as +60 per mil) and refractory carbon

(typically < -140 per mil) with Pannychia ind. #1 (gut sediment ∆14C = -65 to –37 per mil)

ingesting a much greater proportion of the labile carbon than Pannychia ind. #2 (gut sediment

∆14C = -119 to –144 per mil). The analytical error in the 14C measurement is typically 3-8 per

mil, which is about the size of the symbol used in the figure.

14
Figure 2 C data from west Antarctic Peninsula shelf samples. As was the case in the Santa

Catalina Basin samples, the Antarctic animal tissues were enriched in 14C activity relative to the

organic carbon in surface sediments. Particle selection (i.e., the activity difference between the

surface sediments and the foregut contents) was greatest in the holothurian Peniagone sp. and

echiuran worm, whereas assimilative selection processes (i.e., the activity difference between the

foregut contents and the animal tissue) were most evident in the Molpadia sp., the urchin, and

the holothurians, Protelpidia sp. and Bathyplotes sp. The vertical placement of the faunal data

15
within each season was based on the order of increasing 14C enrichment of the gut contents

relative to the surface phytodetritus. The gut sediments in the less particle selective deposit

feeders (i.e., Molpadia sp., Protelpidia sp., and Bathyplotes sp.) were generally more depleted in
14
C activity during the austral winter season (a period of low primary production) than during the

summer, suggesting a decrease in the lability of the organic material available to the benthos

during the winter.

234
Figure 3 Th activities in gut sediments from 6 species of deposit-feeding megafauna (n=48)

collected on the west Antarctic Peninsula continental shelf during 5 cruises between November
234
1999 and March 2001. Th (24 day half life) is scavenged from the water column by settling

particles, which results in high activities in particle trap material (200-975 dpm g-1 in near-

bottom Antarctic shelf particle traps as compared to 10-200 dpm g-1 in surface sediments). The

highest activity gut samples indicate the fauna that are the most particle selective in their feeding

strategies. The numbers shown above the data bars indicate the average activity of all the gut

samples for that species, the standard deviation and the corresponding number of analyses. The

sequence in 234Th gut activities matched and corroborated the sequence of particle selectivity

determined from the 14Corg abundances in the Antarctic surface sediments and gut content

samples presented in Figure 2.

16
Figure 1:

Radiocarbon Data from Santa Catalina Basin

HG = Hindgut Gut Sediment

MG = Midgut Floc Sediment
FG = Foregut Animal Tissue
Local Kelp
Particle Trap
Particle Traps
Local Kelp

Other Benthos

Other Pannychia sp.

HG,FG,MG Pannychia sp. Ind. #2
Pannychia sp . Ind. #1 MG,HG FG
Surface Sediment Floc
Particle Assim.
Selection Select.
Pan. #1 Pan. #1

-200 -150 -100 -50 0 50 100

14
∆ C Activity (per mil)

17
Figure 2:

Radiocarbon Data from the W. Antarctic Peninsula Shelf

HG = Hindgut Winter Data FBIII Gut Sediment-Summer

MG = Midgut Gut Sediment-Winter
FG = Foregut
Animal Tissue
WG = Whole gut Surface Plankton FBIII
Particle Trap Particle Trap
Peniagone sp. WG Surface Plankton
HG FG MG Bathyplotes sp .
Surface Sediment
HG FG Protelpidia sp.
HG FG Molpadia sp.

Surface Plankton FBV Surface Plankton FBII

Particle Trap
WG echiuran
FG,HG WG Peniagone sp.
Bathyplotes sp.
HG,FG Protelpidia sp.
WG urchin
HG FG Molpadia
Surface Sediment Phytodetritus

Summer Data FBV

-250 -225 -200 -175 -150 -125 -100 -75 -50 -25
14
∆ C Activity (per mil)

18
Figure 3.

234
Th Activities in Animal Gut Samples Collected
from the Antarctic Shelf

188+/-98;n=8 149+/-78;n=4 Nov. 1999 Mar. 2000

234 300
Th Jun. 2000 Oct. 2000
114+/-60;n=8 Feb. 2001
Activity 250
in Gut 101+/-44;n=11
200
Samples
36+/-62;n=9
(dpm/g) 150
40+/-28;n=8
100
50
0
1 2 3 4 5 6
Peniagone Echiuran Bathyplotes Protelpidia Urchins Molpadia
sp. (Holoth.) (Worm) sp. (Holoth.) sp .(Holoth.) sp .(Holoth.)
Most Selective Least Selective

19