NAPC 2001
June 26 - July 1 2001 Berkeley, California
Abstracts Ar - Ba
(5/17/01)
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LOSS OF LOCAL ANGIOSPERM SPECIES RICHNESS
PRECEDES THE CRETACEOUS/TERTIARY BOUNDARY:
PALYNOLOGICAL EVIDENCE FOR INCREASED HABITAT HETEROGENEITY
ARENS, Nan Crystal, and Anna Thompson, Dept. of Integrative Biology,
University of California, Berkeley, CA, USA; and A.H. Jahren, Dept. of
Earth and Planetary Sciences, Johns Hopkins University, Baltimore, MD,
USA
Understanding the biological context in which the terminal
Cretaceous bolide impact occurred is important to explaining its biotic
effects. We report palynological data from Russell Basin, McCone County,
northeastern Montana. In this section, the Cretaceous/Tertiary (K/T) boundary
is bracketed biostratigraphically, and diagnosed by a 2 per mil
carbon isotope excursion in terrestrial organic carbon. We recognized
a decline in palynological species richness (the number of distinct forms)
beginning approximately 3.5 m below the Hell Creek-Fort Union formational
contact. Species richness declined from 85 species 3.5 m below the formational
contact to 10 species at about 1 m. Most of the species lost were
triaperturate forms, produced by dicot angiosperms. Relatively low species
richness floras (2634 pollen taxa) persisted across the K/T boundary.
Only about 10 percent of palynospecies present in the uppermost 1 m of
Cretaceous sediment had last occurrences at the K/T boundary. Similar
patterns of local species richness loss in the uppermost meters of the
Cretaceous have been reported throughout Canada. However, these do not
represent pre-K/T boundary extinctions; instead taxa lost at one locality
commonly persist at another. This suggests that the cosmopolitan late
Cretaceous flora was becoming regionally heterogeneous during the interval
represented by the last 3 m of the Cretaceous.
DISTRIBUTION, DIVERSITY, AND LIGHT ISOTOPE SIGNAL
IN BENTHIC FORAMINIFERA ON A SEAMOUNT NEAR THE EAST PACIFIC RISE; EFFECTS
OF THE EAST PACIFIC OXYGEN MINIMUM ZONE
ARNOLD, Anthony J., William C. Parker, and Xiaoli Liu, Dept. of Geological
Sciences, Florida State University, Tallahassee, FL, USA
Seamount 7 is about 30 km from the axis of the East Pacific
Rise at 102° 30' W, 13° 25' N at depths ranging from 730 to
3,353 m below MSL; at its summit Seamount 7 penetrates into the East Pacific
oxygen minimum zone (OMZ) thus affording an unusual opportunity to investigate
the effects of oxygen deprivation on benthic foraminiferal communities
in an open marine environment. Bottom water oxygen concentrations vary
from near-microxic (about 0.1 ml/l) at the top of Seamount 7 to dysoxic
(0.1 to 1 ml/l) at about 1,100 m below the summit. At greater depths,
the bottom waters would be considered oxic. The likely presence of the
OMZ at Seamount 7 (as well as at two other seamounts in the area) was
reported in 1987 based on foraminiferal diversity, test ultrastructural
evidence, and relative species abundance data consistent with other reports
of foraminiferal response to oxygen deprivation. Later studies focused
on macrofaunal and physio-chemical characterization of Seamount 7 with
special attention to the effects of oxygen concentrations, and provided
an excellent background for understanding foraminiferal distribution patterns.
The present study focuses exclusively on Seamount 7 in the context of
that background data and using a new and more comprehensive suite of samples
with the following goals in mind: (1) to test the hypothesis that stable
light isotope signals, particularly 12C/13C, should
reflect the presence of an oxygen minimum, and (2) to refine and quantify
our understanding of the effects of oxygen deprivation on the foraminiferal
populations in this area with respect to distribution, diversity, abundance,
and test ultrastructure. Shannon-Weaver diversity values are significantly
lower within the OMZ. This is primarily caused by the high dominance of
cassidulinids, while Ehrenbergina spp showed high abundances and unusually
thin shells. Relatively elevated frequencies of bolivinids and uvigerinids
were also noteworthy in the OMZ. Elevated diversities were characteristic
of the ecotone separating the OMZ from oxic conditions.
NATICID GASTROPOD DRILLHOLES: AN INTEGRATIVE
APPROACH INVOLVING RE-ANALYSIS OF PUBLISHED DATA, EXPERIMENTAL TESTS,
AND INTERNAL BIOLOGY
ARONOWSKY, Audrey, Dept. of Integrative Biology and Museum of Paleontology,
University of California, Berkeley, CA, USA
Naticids are infaunal predatory gastropods that drill
holes in the shells of their molluscan prey. Because these drillholes
are beveled, they have two diameters, an inner-boring diameter (IBD) and
an outer boring diameter (OBD). In a seminal paper, Kitchell et al. (1981)
reported a positive correlation of OBD with the shell length of the drilling
naticid. This correlation is thought to represent a relationship between
the size of the predator and its inflated accessory boring organ (ABO)
diameter. Re-analysis of the Kitchell et al. (1981) data suggests that
this relationship is more tenuous than previously thought. For example,
an OBD value of 3.25 mm can yield shell length values ranging from 2952
mm. In addition, the Kitchell et al. (1981) data are derived from the
analysis of a single naticid species, Polinices duplicatus.
Experimental studies on other naticid species will test
whether the shell length:OBD correlation is valid for other taxa. Laboratory
studies will maintain individual naticids with prey of varying morphology
to examine the range of drillhole geometries produced by an individual
predator. This technique will provide a more rigorous investigation into
the relationship between naticid size and drillholes produced. Subsequent
dissection of these same predators will permit the investigation of the
ABO:OBD relationship. Currently, categorization of naticid drillholes
(fossil or recent) below the family level is problematic and will only
be elucidated through analysis of many naticid species.
ACTIVITY OF PREDATORY GASTROPODS ON THE SHELLS
OF LATE-OLIGOCENE (EGERIAN) MOLLUSCS (WIND BRICKYARD, EGER, HUNGARY)
ARPAD, David, Karoly Eszterhazy College, Dept. of Geography, Eger, Hungary
The clay-pit of Wind Brickyard at Eger is the stratotype
of the Egerian Stage. Sequence of the profile is: glauconitic sandstone;
molluscan clay; marine silty sandstone; marine limonitic sandstone; alternation
of coarse sand and carbonaceous clay. These formations are rich in molluscan
fossils. 22,065 specimens were examined, belonging to 189 species. Shells
of bivalves, gastropods and scaphopods showed traces of predatory gastropods
belonging to the families Naticidae and Muricidae. Mode of life, feeding
habits of the prey molluscs, and distribution of successful, unsuccessful
and unfinished borings were investigated and compared. The molluscs examined
have been found in the following layers: molluscan clay, marine silty
sandstone, marine limonitic sandstone, alternating coarse sand and carbonaceous
clay. Naticid boreholes were examined on the shells of molluscs collected
from three different layers of the exposure. Traces of naticid predation
occurred on the shells of bivalves, gastropods and scaphopods. Mainly
gastropods were attacked by naticids, producing 998 boreholes on 944 shells,
with successful borings the most frequent. The rate of bored specimens
is 4.44% in the samples of the Molluscan Clay. Increasing number of naticid
borings were observed in molluscs of the other layers. Multiple naticid
borings and "cannibalism" have also have been found. Suspension
feeders and scavengers are dominant among the prey species of naticids.
Occurence of muricid drillholes on the shells of molluscs collected from
four different layers of Wind Brickyard was examined. Traces of muricids
were found on the shells of bivalves and gastropods, with 220 borings
observed on 201 shells. Gastropods were the most common prey; successful
borings were dominant. The number of boreholes was low on the molluscs
collected from the Molluscan Clay with increased numbers in the upper
layers. The only layer where the number of unfinished borings was higher
than the two other boring types was the uppermost layer. Suspension feeders
and scavangers were the main prey items of muricids.
THE ENTOBIA ICHNOGENUS IN HUNGARIAN TERTIARY
FORMATIONS
ARPAD, David, Monika Brecz, and Judit Horvath, Karoly Eszterhazy College,
Dept. of Geography, Eger, Hungary
Occurrence and distribution of Entobia ichnospecies
at eight Hungarian Tertiary localities have been examined. The age of
the localities are middle Eocene (Lutetian), early Miocene (Egerian, Ottnangian,
Carpathian) and middle Miocene (Badenian). Oyster shells, gastropod tests
and abrasion pebbles served as firm substrate for the Clionid larvae.
One hundred twenty epoxy-casts have been made from the collected material
(515 from each locality). From these epoxy casts, seven Entobia
ichnospecies have been identified. They include the following: Entobia
cateniformis Bromley et D'Alessandro, E. geometrica Bromley
et D'Alessandro, E. laquea Bromley et D'Alessandro, E. megastoma
(Fischer), E. ovula Bromley et D'Alessandro, E. paradoxa
(Fischer), and E. retiformis (Stephenson). The most frequent ichnospecies
are E. geometrica, E. cateniformis and E. megastoma.
Idiomorphic forms are rare in the oyster valves. Depth of the trace fossils
is shallow. Clionid borings also occur in the inner side of the valves
in most cases. From our observations, we found no correlation between
the thickness of the oyster valves and the depth of the Entobian borings.
"D+" growth phase occur only in the Entobia ichnospecies
observed in the tests of gastropods. Growth phases from "A"
to "D" occur in the case of the Entobians of the abrasion pebbles.
Forms are idiomorphic in each case. Based on our observations, Clionid
sponges played a significant role in the destruction of shelly substrates
and calcareous rocks during the Tertiary of Hungary.
OCCURENCE AND DISTRIBUTION OF GASTROCHAENOLITES
ICHNOGENUS IN MIDDLE MIOCENE (CARPATHIAN) AGE ABRASION PEBBLES OF TWO
LOCALITIES
A COMPARISON (NAGYVISNYO & DEDESTAPOLCSANY BÜKK MOUNTAINS, HUNGARY)
ARPAD, David, and Rozalia Fodor, Karoly Eszterhazy College, Dept. of
Geography, Eger, Hungary
Abrasion pebbles from two localities (Nagyvisnyo, abandoned
limestone quarry and Dedestaapolcsany, pebble quarry) have been examined
for the presence, abundance and distribution of Gastrochaenolites
ichnospecies. The Gastrochaenolites ichnogenus is formed by boring
bivalves in calcareous substrates. Their characteristic club-shaped trace
fossils are dwelling structures (domichnia). The age of the black Permian
limestone and dolomite pebbles is middle Miocene (Carpathian) and belongs
to the Egyhazasgerge Formation. Epoxy casts of pebbles from both localities
were made: 36 from Nagyvisnyo and 43 from Dedestapolcsany. Among the number
of observed bioerosional traces of boring bivalves, these could be attributed
to the following taxa: G. lapidicus, G. cor, G. cluniformis, G. orbicularis,
G. torpedo and G. turbinatus. The frequency of G. lapidicus
and G. torpedo is significant at both localities. The high number
of G. orbicularis in the pebbles collected at Dedestapolcsany is
also remarkable. Differences in the length of G. lapidicus and
G. torpedo at the two localities is attributed to paleoenvironmental
alternations; in the case of Dedestapolcsany locality the fossil rocky
shore was richer in nips and uneven surfaces.
TWO FLORAS IN THE UPPER JOHN DAY FORMATION NEAR
MADRAS, CENTRAL OREGON
ASHWILL, Melvin S., Madras, OR, USA
Two fossil floras in the upper John Day Formation, one
well collected, the other reconnaissance collected, add to the understanding
of this poorly documented gap in the paleobotanical record of the early
Miocene in central Oregon. The data presented tend to corroborate previous
observations about the drying and cooling climatic change that took place
there between early Oligocene and late Miocene times. The gradual replacement
in the niche once held by the dominant Metasequoia ("dawn
redwood") by Taxodium (swamp cypress) is relected in these
floras.
INTERPRETATION OF BIOLOGICAL AND ENVIRONMENTAL
CHANGES ACROSS THE NEOPROTEROZOIC-
CAMBRIAN BOUNDARY: DEVELOPING TOOLS FOR
PREDICTING THE OCCURRENCE OF BURGESS SHALE-TYPE DEPOSITS
BABCOCK, Loren E., Dept. of Geological Sciences, The Ohio State University,
Columbus, OH, USA
The fossil record of biotic change from the late Neoproterozoic
through the Middle Cambrian resulted from an interplay of biological and
environmental factors. By the Early Cambrian, biological conditions underway
in shallow shelf seas included rapid diversification, experimentation
with new body plans, and a shifting of ecological setting away from mat-dominated
communities to more burrowed sediment surfaces. High eustatic sea level,
high tidal ranges, areas of rapid sedimentation, and interfaces between
conditions of contrasting seawater chemistry afforded opportunities for
the exceptional preservation of nonmineralizing organisms at a rate exceeding
that for any other interval of the Phanerozoic.
Understanding the conditions under which exceptional
preservation occurred during the Early and Middle Cambrian provides a
framework for exploring for previously unknown sites of exceptional preservation.
For the most part, Burgess Shale-type deposits were developed in tropical
to subtropical paleolatitudes where sedimentation was relatively rapid,
and commonly episodic. Exceptional preservation seems to have occurred
most commonly in late phases of transgressive systems or in highstand
systems tracts. Such preservation is typically associated with thin bedding
of clay- or silt-sized grains. Rhythmic bedding patterns in some localities
suggest tidally influenced deposition. Oxic-anoxic, saline-hyposaline,
and potentially saline-hypersaline, conditions are inferred to have inhibited
scavengers, microbial biodegraders, and bioturbators, thus providing sufficient
time for burial prior to the occurrence of substantial breakdown of organic
remains. Following burial, geochemical conditions, including establishment
of dysoxic or reducing microenvironments, would have promoted exceptional
preservation of nonmineralized remains.
DATABASES FOR RECENT MAMMALS AND MAMMALIAN FAUNAS
BADGLEY, Catherine, Museum of Paleontology, University of Michigan, Ann
Arbor, MI, USA; and David L. Fox, Dept. of Earth Sciences, University
of California, Santa Cruz, CA, USA
Databases for modern mammalian faunas were created to
address questions in community ecology, biogeography, and paleoecology.
Mammals are well documented in terms of taxonomy, ecology, and geographic
distribution, and thus are well suited to coverage in databases and synoptic
analyses. We constructed three kinds of database. One is a species file
that records taxonomic and ecological information for individual species.
Individual species have multiple records when their ecological attributes
show high geographic variation, as in species with clinal variation in
body size. Second is a locality file based on sites where comprehensive
faunal surveys occurred. This database records geographic, ecological,
and climatic information about each locality. The third database contains
quadrats from a grid system that covers continental regions. For each
quadrat, we recorded latitude, longitude, and topographic and climatic
data. The species database is linked to the site databases through a relational
file that expresses species occurrences at particular sites. We used these
databases to quantify the relationship between environmental variables
and ecological diversity of modern mammalian faunas. For example, we studied
the ecological biogeography of all 721 extant species of North American
mammals. Five climatic and topographic variables explain 88% of the geographic
variation in species density, including the latitudinal and longitudinal
gradients in species density. Five environmental variables explain 76%
of the variation in mammalian ecological diversity, as expressed by size
and trophic structure. Evaluating how much these patterns reflect ecological
sorting of species ranges since the last deglaciation versus geographic
variation in speciation and extinction will require linking our research
goals and databases with those focusing on Quaternary and Neogene mammals,
vegetation, and environments. We also see opportunities for productive
links with databases serving biogeographic or conservation goals for extant
organisms and environments.
AN ANALYSIS OF PALYNOMORPHS OF THE AGUJA FORMATION
(CAMPANIAN), BIG BEND NATIONAL PARK, BREWSTER COUNTY, TEXAS
BAGHAI-RIDING, Nina L., Dept. of Biological and Environmental Sciences,
Delta State University, Cleveland, MS, USA
Exposures of the Aguja Formation (8474.5 m.y.a.)
at Big Bend National Park contain deltaic sequences that were deposited
on the southwestern margins of the Western Interior Seaway. Near-shore
marine and paralic sandstone units interbed lignite, carbonaceous shales
and mudstones. Well-preserved assemblages of palynomorphs provide greater
resolution of the two-progradational deltaic cycles than are reflected
by sedimentological and stratigraphic data. The microfossil flora also
provides an accurate assessment of subtropical to warm temperate paleoclimates
and concurrent plant communities that once prevailed in this region.
Palynomorph samples, collected and processed from six
measured and correlated stratigraphic sections from the northwestern and
northeastern part of Big Bend, include a diverse assortment of freshwater
protists, dinoflagellates, fungi and vascular and nonvascular plant taxa.
Quantitative counts and analysis of 54 samples revealed 134 various form
genera: 28 dinoflagellates and freshwater algae, 16 fungi, 37 ferns and
mosses, 13 gymnosperms and 40 angiosperms. The lower shale member, which
accumulated largely in coastal marshes and freshwater swamps, possesses
a more diverse array of fungi, ferns, mosses and angiosperms than the
upper two shale units. The thin, middle shale unit contains a larger assortment
of dinoflagellates at its base that reflects a marginal marine influence.
The upper shale unit, which was deposited in coastal floodplains, freshwater
swamps and continental floodplains, contains many tricolporate and tricolpate
pollen grains, not found in the other horizons. Common genera are Deflandrea,
Palaeohystrichophora (dinoflagellates), Schizosporis (freshwater
algae), Cyathidites, Deltoidospora, Gleicheniidites (ferns), Cycadopites,
Inaperturopollenites (gymnosperms), and Liliacidites, Momipites,
Plicapollis, and Proteacidites (angiosperms).
WALKER AND BAMBACH 1971 REVISITED: AT LAST WE
ARE RELEVANT
BAMBACH, Richard K., Dept. of Geological Sciences, Virginia Tech, Blacksburg,
VA, USA
Walker and Bambach 1971 (a GSA meeting discussion paper
in the days before poster sessions) was inspired by the difficulty in
communication between biologists and paleontologists that characterized
the Penrose Conference on marine ecology and paleoecology held in December,
1970. The biologists were skeptical that ecologically useful information
could be obtained from accumulations of non-contemporaneous specimens,
yet that is the stuff of fossil assemblages.
Times have changed. Time-averaged assemblages are now
recognized as having impressive environmental fidelity. Because time-averaging
reduces the variability that short-term patchiness and response to disturbance
generates, time-averaged assemblages often resolve environmental gradients
in greater detail than can be detected with sedimentological analysis
alone. Since the fossils preserved in a time-averaged deposit lived (and
died) during the interval after the underlying bed was deposited and before
the end to disturbance and deposition marked by the bed in which the fossils
are preserved, the fossils in a time-averaged deposit actually record
information from the time otherwise concentrated in bedding-plane diastems.
Dating of individual shells by amino-acid racimization and calibration
from radiocarbon dating permits segregation of different age shells from
time-averaged deposits in the sub-Recent and Recent. This offers the opportunity
of monitoring isotopic patterns and following environmental change even
during intervals of time-averaging. Environmental patterns on century
to millennial scales may become available from deposits that mix shells
from disparate times together in such time-averaged, reworked and redeposited
assemblages. Time-averaged but little mixed assemblages are now recognized
as having great utility in understanding biologically relevant phenomena
in the past.
A NEW FAUNA OF VERY MAMMAL-LIKE CYNODONTS FROM
THE LATE TRIASSIC OF BRAZIL
BARBERENA, M.C, and J.F. Bonaparte, Federal University of Rio Grande
do Sul, Porto Alegre, Brazil; A.W. Crompton, Museum of Comparative Zoology,
Harvard University, Cambridge, MA, USA; J.A. Hopson, Dept. of Organismal
Biology and Anatomy, University of Chicago, Chicago, IL, USA; and C. Schultz
and R. Rubert, Instituto de Geociencias, Universidad Federal de Rio Grande
do Sul, Porto Alegre, Brazil
Very small cynodont therapsids of the Late Triassic have
long been assumed to have given rise to mammals. However, with the exception
of Therioherpeton cargnini, described from an incomplete skull
from the Santa Maria Formation of Brazil and isolated teeth of uncertain
affinities from Europe and North America, such small cynodonts have been
unknown. A new fauna of extremely mammal-like cynodonts has recently been
discovered in the Caturrita "Formation" in Rio Grande do Sul,
Brazil. This fauna is associated with sphenodontids and procolophonids
and is considered to be of early Norian Age.
Several skulls with associated lower jaws of a primitive
ictidosaur, Riograndia quaibaensis, recently described by Bonaparte
et al., and a collection of partial skulls, lower jaws and postcranial
elements representing very small mammal-like cynodonts are briefly discussed.
Riograndia appears to be related to the classic
South African ictidosaurs (the trithelodontids Pachygenelus, Diarthrognathus
and Trithelodon), but it is both more primitive in cranial anatomy
and divergently specialized in its dentition.
The remaining cynodonts are much smaller, with more slender
delicate skulls and lower jaws. Their postcanine teeth resemble those
of Morganucodon and other basal mammalian forms, but they lack
certain diagnostic features seen in Early Jurassic mammaliformes. These
Late Triassic non-mammalian cynodonts appear to be more closely related
to mammals than either the trithelodontids or tritylodontids.
The entire cynodont fauna is currently being studied
with cooperation between the Federal University of Rio Grande do Sul and
the Natural Science Museum of Buenos Aires, while the species of "ictidosaurs"
are being studied in cooperation with the paleontologists from the University
of Chicago and Harvard.
MIOMAP: A RELATIONAL AND SPATIAL DATABASE FOR
RESEARCH ON EVOLUTION, ECOLOGY, AND BIOGEOGRAPHY OF MIOCENE MAMMALS
BARNOSKY, Anthony D., and Marc A. Carrasco, Museum of Paleontology and
Dept. of Integrative Biology, University of California, Berkeley, CA,
USA
The Miocene Mammal Mapping Project (MIOMAP) has the primary
goal of casting light on the question: How do large-scale physical-environmental
perturbations affect the evolution and biogeography of mammals? The intent
is to apply the fossil record to help delineate what is natural for the
mammalian component of ecosystems to assess as accurately as possible
the environmental effects of human induced global change. MIOMAP is a
relational database in PARADOX 9 designed for incorporation into a GIS
analytical tool (ARC/INFO 8.0). It includes all published Miocene mammal
species of the United States, and as time and resources allow, will also
include well-identified, unpublished specimens represented by voucher
specimens in museums. Information associated with each species occurrence
includes latitude, longitude, relevant taxonomy, geologic age, depositional
environment, taphonomic attributes, bibliographic information, and numbers
of specimens, among other relevant data. The data structure parallels
that for the FAUNMAP project, with which complete compatibility is envisioned.
A later phase of the project also calls for exploring interfaces that
would easily link MIOMAP with other relevant datasets. The immediate research
application of MIOMAP is to test hypotheses about how global warming and
a tectonic event ~17 Ma ago affected faunal turnover, species richness,
and biogeographic patterns in the western US. Analyses so far have focused
on comparisons of the Pacific Northwest, Northern Rockies, and northern
Great Plains. Preliminary results indicate that alpha diversity and faunal
turnover rates in the northern Rockies may have increased as a result
of the environmental perturbations and that beta diversity may have remained
the same or decreased. However, the results highly depend on analytical
method. In addition, the three areas seem to have retained biogeographic
integrity throughout the Miocene intervals that were studied; modern biogeographic
patterns may therefore be deeply rooted in time.
COST-BENEFIT ANALYSIS AS A GUIDE TO THE ECOLOGY
OF DRILLING PLATYCERATIDS
BAUMILLER, Tomasz K., Museum of Paleontology, University of Michigan,
Ann Arbor, MI, USA
Boreholes in Paleozoic crinoids and blastoids have been
hypothesized as parasitic based on the presence of: (1) multiple and healed
borings, which indicate that drilling was not fatal, (2) attachment scars,
which imply a long-term association between host and parasite. It has
also been argued that these boreholes were made by platyceratids because
examples of platyceratid infested blastoids and crinoids are known, and
one example of a platyceratid positioned over a borehole on a crinoid
tegmen has been reported.
To further explore the platyceratid parasitic drilling
hypothesis, a cost-benefit analysis was performed comparing the quality
of various crinoids as hosts and as edible prey. Platyceratid energetics
were modeled on the marine snail Polinices, for which experimental
data on drilling and metabolic rates exist. The value of crinoids as prey
was calculated using the empirical relationship between calyx soft tissues
and calyx size, and data on caloric content of soft tissues in extant
crinoids. The value of crinoids as hosts was evaluated assuming snails
"stole" unused captured nutrients directly, or through coprophagy.
Crinoid nutrient capture was modeled for a range of filter morphologies,
concentrations of particles in seawater, and current velocities, using
previously developed models for passive suspension feeders.
Results indicate that under realistic environmental conditions
pelmatozoans with high solidity filters (pinnulate or highly branched)
(1) could capture sufficient nutrients to host kleptoparasitic snails,
(2) provided a greater energetic return as hosts than as prey items, and
(3) provided a greater energetic return as hosts than would be available
for snails preying on contemporaneous brachiopods. This may partly explain
the host preferences of platyceratids (associated only with pelmatozoans
with high-solidity filters) and the apparent failure of these snails to
exploit the predatory potential of drilling.
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