BEN # 282

Adolf Ceska aceska at victoria.tc.ca
Wed Feb 20 09:33:08 EST 2002


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No. 282                              February 20, 2002

aceska at victoria.tc.ca                Victoria, B.C.
-----------------------------------------------------------
 Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2
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               This issue of BEN is dedicated to

                           PHYTOLITHS

       those small specks that botanists tend to overlook


PHYTOLITH LITERATURE REVIEW
From: Terry B. Ball [tbball at reled.byu.edu]

Attached is a brief intro to Phytoliths taken form the following
publication:

Ball,  T.B.,  and  J. D. Brotherson. 1992. The effect of varying
   environmental  conditions  on  phytolith  morphology  in  two
   species  of  grass  (_Bouteloua  curtipendula_  and  _Panicum
   virgatum_). Scanning Electron Microscopy 6:1163-1182.

Other more recent publications in which I discuss similar topics
include:

Ball, T.B., J.D. Brotherson, and J.S. Gardner. 1996. Identifying
   phytoliths produced by  the  inflorescence  bracts  of  three
   species  of  wheat  (_Triticum  monoccocum_ L., _T. dicoccon_
   Schrank., and _T. aestivum_ L.) using computer-assisted image
   and statistical analyses. _Journal of Archaeological Science_
   23:619-632.
and
Ball, T.B., J.D. Brotherson, and J.S. Gardner. 1993. A typologic
   and morphometric  study  of  phytoliths  from  einkorn  wheat
   (_Triticum  monococcum_  L.).  _Canadian  Journal  of Botany_
   71:1182-1192.

I also have a phytolith webpage at
http://reled.byu.edu/ascrip/tball
[this link did not work on 2002/02/20]

Monosilicic acid in the soil, created  from  the  weathering  of
rocks  and  the  dissolution  of biologically deposited SiO2, is
taken  up  by  plant  roots.  Following  uptake  the   acid   is
transported  to  various plant organs, where, in many taxa, some
of it polymerizes to form  solid  silica  deposits  at  specific
intracellular  and  extracellular locations (Jones and Handreck,
1967; Raven, 1983; Sangster,  1970).  These  solid  deposits  of
SiO2,  as  well  as  deposits containing calcium compounds, have
been given  the  name  "phytolith",  literally  meaning  "plant-
rocks."   Many  plants  produce  phytoliths  with  morphological
characteristics  that  appear  unique  to  a  given   taxon,   a
phenomenon giving them taxonomic significance.

There  has  been  considerable  interest  in phytolith research.
Phytolith formation and deposition in various cereal grasses has
been well documented (Blackman, 1968, 1969; Blackman and  Parry,
1968; Hayward and Parry, 1973; Hodson and Sangster, 1989; Hutton
and  Norrish,  1974;  Jones  and Handreck, 1965; Kaufman et al.,
1972; Soni and Parry, 1973). The role  of  phytoliths  in  plant
resistance  to  disease  and  insects  has been investigated (De
Silva and Hillis, 1980; Djamin and Pathak, 1967; Hanifa et  al.,
1974;  Jones  and Handreck, 1967; Kunoh and Ishizaki, 1975; Lan-
ning, 1966), as well as the detrimental effects phytoliths  have
on  herbivores  and  humans  (Baker,  1961,  Baker et al., 1959;
Bezeau et al., 1966; Forman and Sauer,  1962;  Harbers  et  al.,
1981;  O'Neill  et  al.,  1982; Parry and Hodson, 1982; Bhatt et
al., 1984). Phytolith research has  proved  highly  valuable  to
archaeobotanists. Because phytoliths are siliceous, when a plant
dies,  even if it is burned, buried, or ingested, its phytoliths
persist and maintain their morphological integrity,  becoming  a
microfossil  of  that  plant.  Microfossil  phytoliths have been
collected by archaeobotanists from such diverse environments  as
paleosols  exposed  by  erosion  or  excavation  (Piperno, 1983,
1988), ceramics and bricks made from clay upon which  vegetation
once  grew,  or to which plant fibers were added (Rands and Bar-
gielski, paper presented at the 1986 meeting of the Society  for
American  Archaeology) tooth tartar and coproliths of herbivores
(Bryant, 1974; Armitage, 1975), and the surface of  stone  tools
used  to  process  plants  and/or  plant  parts (Kamminga, 1979;
Anderson, 1980).

Once collected and analyzed, microfossil phytoliths can  provide
researchers with significant information and insights. Microfos-
sil phytoliths have been used for the reconstruction of paleoen-
vironments  (Fisher  et  al., 1987; Lewis, 1981; Robinson, 1979;
Rovner, 1971; Twiss, 1987), as indicators of ancient  industrial
and  agricultural  practices (Liebowitz and Folk, 1980; Piperno,
1984; Rosen, 1992; Rosen, 1999), and for tracing the origins and
developments of cultigens  (Piperno,  1988).  Rovner  (1983)  in
reviewing  the  value  and  advances of phytolith research, sug-
gested that it has the potential to become a second  palynology.
Pearsall  (1989)  and  Piperno  (1988)  point out that phytolith
analysis is especially valuable to archaeobotanists at sites  of
study were other plant remains are absent. They further indicate
that  when  phytoliths  are used in conjunction with other plant
remains, they add precision and support for any  interpretations
made.

References:

Anderson,  P.C. 1980. A testimony of prehistoric tasks: diagnos-
   tic residues on stone tool working edges. _World archaeology_
   12: 181-194.
Armitage, P.L. 1975. The extraction and identification  of  opal
   phytoliths  from  the  teeth  of  ungulates.  _Journal of Ar-
   chaeological Science_ 2: 187-197.
Baker, G. 1961. _Opal phytoliths and adventitious  mineral  par-
   ticles in wheat dust._ Commonwealth Scientific and Industrial
   Research  Organization,  Australia; Mineralgraphic Investiga-
   tions Technical Paper No. 4, 3-l2.
Baker, G., L.H.P. Jones, & I.D. Wardrop. 1959. The cause of wear
   in sheep's teeth. _Nature_ 184: 1583-4.
Bezeau L.M., A. Johnson, & S. Smoliak. 1966. Silica and  protein
   content  of mixed prairie and fescue grassland vegetation and
   its relationship to the  incidence  of  silica  urolithiasis.
   _Canadian Journal of Plant Science_ 46: 625-631.
Bhatt,  T.S., M.M. Coombs, & C.H. O'Neill. 1984. Biogenic silica
   fibre promotes carcinogenesis in mouse  skin.  _International
   Journal of Cancer_ 34: 519-528.
Blackman,  E.  1968.  The  pattern  and  sequence of opaline silica
   deposition in rye (_Secale cereale_ L.). _Annals  of  Botany_
   32: 207-218.
Blackman, E. 1969. Observations on the development of the silica
   cells  of  the  leaf  sheath  of wheat (_Triticum aestivum_).
   _Canadian Journal of Botany_ 47: 827-838.
Blackman, E. 1971. Opaline silica bodies in the range grasses of
   southern Alberta. _Canadian Journal of Botany_ 49: 769-81.
Blackman, E. & D.W. Parry. 1968. Opaline  silica  deposition  in
   rye (_Secale cereale_ L.). _Annals of Botany_ 32: 199-206.
Bozarth,  S.R.  1987.  Diagnostic  opal phytoliths from rinds of
   selected _Cucurbita_ species. _American Antiquity_  52:  607-
   615.
Brown,  D.  1984.  Prospects  and  limits of a phytolith key for
   grasses  in  the  central  United  States.  _Journal  of  Ar-
   chaeological Science_ 11: 221-243.
Bryant,  V.M.,  Jr.  1974. The role of coprolith analysis in Ar-
   chaeology. _Texas Archaeological Society Bulletin_ 45: 1-28.
De Silva, D.& W.E. Hillis. 1980. The contribution of  silica  to
   the  resistance of wood to marine borers. _Holzforschung_ 34:
   95-97.
Dayanandan, P., P.B. Kaufman, & C.I. Franklin.  1983.  Detection
   of  silica  in plants. _American Journal of Botany_ 70: 1079-
   1084.
Djamin, A. & M.D. Pathak. 1967. Role of silica in resistance  to
   asiatic  rice  borer,  _Chilo suppressalis_ (Walker), in rice
   varieties. _Journal of Economic Entomology_ 60: 347-351.
Fisher, R.F., M.J. Jenkins, & W.F. Fisher. 1987.  Fire  and  the
   prairie-forest mosaic of Devils Tower National Monument. _The
   American Midland Naturalist_ 117: 250-257.
Forman,  S.A.  &  F.  Sauer.  1962. Some changes in the urine of
   sheep fed a hay high in silica. _Canadian Journal  of  Animal
   Science_ 42: 9-17.
Gould,  F.W. & R.B. Shaw. 1983. _Grass Sytematics_, 2nd edition,
   Texas A & M University Press, College Station 226.
Hanifa, A.M., T.R. Subramaniam, & B.W.X. Ponnaiya. 1974. Role of
   silica in resistance  to  the  leaf  roller,  _Cnaphalocrocis
   medinalis_  Guenee,  in rice. _Indian Journal of Experimental
   Biology_ 12: 463-465.
Harbers, L.H., R.J. Raiten, & G.M. Paulsen. 1981.  The  role  of
   plant  epidermal  silica  as  a structural inhibitor of rumen
   microbial digestion in steers.  _Nutrition  Reports  Interna-
   tional_ 24: 1057-1066.
Hayward,  D.M.  & D.W. Parry. 1973. Electron-probe microanalysis
   studies of silica deposition  in  barley  (_Hordeum  sativum_
   L.). _Annals of Botany_ 37: 579-591.
Hodson,  M.J.  &  A.G.  Sangster. 1989. Silica deposition in the
   inflorescence bracts of wheat (_Triticum aestivum_).  II.  X-
   ray   microanalysis   and   backscattered  electron  imaging.
   _Canadian Journal of Botany_ 67: 281-287.
Hutton, J.T. & K. Norrish. 1974. Silicon content of wheat  husks
   in  relation  to  water  transpired.  _Australian  Journal of
   Agricultural Research_ 25: 203-212.
Jones, L.H.P. & K.A. Handreck. 1965. Studies of  Silica  in  the
   oat  plant.  III.  Uptake  of silica from soils by the plant.
   _Plant and Soil_ 23: 79-96.
Jones, L.H.P. & K.A. Handreck. 1967. Silica in soils plants  and
   animals. _Advances in Agronomy_ 19: 107-149.
Kamminga, J. 1979. The nature of use-polish and abrasive smooth-
   ing on stone tools. Pp. 143-157 in : Hayden, B. (ed.) _Lithic
   Use-wear Analysis_, Academic Press, New York.
Kaufman,  P.B.,  S.L.  Soni,  J.D.  Lacroix,  J.J. Rosen, & W.C.
   Bigelow. 1972. Electron-probe microanalysis of silicon in the
   epidermis of rice (_Oryza sativa_  L.)  internodes.  _Planta_
   104: 10-17.
Kunoh,  H.  & H. Ishizaki. 1975. Silicon levels near penetration
   sites of fungi on wheat, barley, cucumber, and morning  glory
   leaves. _Physiological Plant Pathology_ 5: 283-287.
Lanning, F.C. 1966. Barley silica: relation of silicon in barley
   to  disease,  cold, and pest resistance. _Journal of Agricul-
   ture and Food Chemistry_ 14: 636-638.
Lewis, R.O. 1981. Use of opal phytoliths  in  paleoenvironmental
   reconstruction. _Journal of Ethnobiology_ 1: 175-181.
Liebowitz,  H.  & R.L. Folk. 1980. Archaeological geology of Tel
   Yin'am, Galilee, Israel. _Journal of  Field  Archaeology_  7:
   23-42.
Mulholland,  S.C.  &  G.R.  Rapp,  Jr. 1989. Characterization of
   grass  phytoliths  for  archaeological  analysis.  _Materials
   Research Bulletin_ 14: 36-39.
Ollendorf  A.L., S.C. Mulholland, & G. Rapp, Jr. 1988. Phytolith
   analysis as a means of plant identification:  _Arundo  donax_
   and _Phragmites communis_. _Annals of Botany_ 61: 209-214.
O'Neill, C.H., Q. Pan, G. Clarke, F.S. Liu, G. Hodges, M. Ge, P.
   Jordon,  Y.M.  Chang, R. Newman, & & E. Toulson. 1982. Silica
   fragments from millet bran in mucosa surrounding  oesophageal
   tumors in patients in northern China. _The Lancet_ 15(May 29,
   1982): 1202-1206.
Parry, D.W. & F. Smithson. 1964. Types of opaline silica deposi-
   tions  in  the  leaves of British grasses. _Annals of Botany,
   N.S._ 28(109): 169-85.
Parry, D.W. & F. Smithson.  1966.  Opaline  silica  in  the  in-
   floresences  of  some British grasses and cereals. _Annals of
   Botany, N.S._ 30(119): 525-38.
Parry, D.W. & M.J. Hodson.  1982.  Silica  distribution  in  the
   caryopsis   and   inflorescence   bracts  of  foxtail  millet
   (_Setaria italica_) and its  possible  significance  in  car-
   cinogenesis. _Annals of Botan, N.S._ 49: 531-540.
Pearsall, D.M. 1978. Phytolith analysis of archaeological soils:
   evidence   for   maize   cultivation  in  formative  Equador.
   _Science_ 199: 177-178.
Pearsall,  D.M.   1989.   _Paleoethnobotany:   A   Handbook   of
   Procedures._ Academic Press, San Diego.
Piperno,  D.R.  1983.  _The application of phytolith analysis to
   the reconstruction of plant subsistence and  environments  in
   prehistoric Panama._ Ph.D. Dissertation, Temple University.
Piperno,   D.R.   1984.  A  comparison  and  differentiation  of
   phytoliths from maize (_Zea mays_ L.) and wild  grasses:  Use
   of morphological criteria. _American Antiquity_ 49: 361-383.
Piperno,  D.R.  1985.  Phytolith  analysis  and  tropical paleo-
   ecology: production and taxonomic significance  of  siliceous
   forms  in  New  World  plant  domesticates  and wild species.
   _Review of Paleobotany and Palynology_ 45: 185-228.
Piperno, D.R. 1988. _Phytoliths analysis: An archaeological  and
   geological perspective._ Academic Press, San Diego.
Rapp,   G.R.,   Jr.   1986.   Morphological   classification  of
   phytoliths, Pp.  33-35  in:  Rovner,  I.  (ed.)  _Plant  opal
   phytolith  analysis in archaeology and paleoecology, Proceed-
   ings of the 1984 Phytolith Research Workshop, North  Carolina
   State University, Raleigh, Occasional Papers No. 1 Raleigh.
Raven,  J.A.  1983.  The  transport  and  function of silicon in
   plants. _Biological Reviews of  the  Cambridge  Philosophical
   Society_ 58: 179-207.
Robinson,  R.L.  1979.  Biosilica  analysis:  paleoenvironmental
   reconstruction of 41 LL 254. Appendix III  in:  Assad,  C.  &
   D.R. Porter. _An Intensive Archaeological Survey of Enchanted
   Rock  State Natural Area._ Center for Archaeological Research
   Survey Report 84, San Antonio.
Rosen, A.M. 1992. Preliminary identification of silica skeletons
   from near eastern archaeological  sites:  an  anatomical  ap-
   proach.  Pp.  129-147 in: Mulholland S. & G. Rapp, Jr. (eds.)
   _Phytolith systematics: Emerging issues._, Plenum Press,  New
   York.
Rosen,  A, 1999. Phytoliths as indicators of prehistoric irriga-
   tion farming, Pp. 193-198 in: Anderson, P.C.  (ed.)  _Prehis-
   tory   of  Agriculture:  New  Experimental  and  Ethnographic
   Approaches._  193-198.  UCLA,Institute  of  Archaeology,  Los
   Angeles.
Rovner,  I.  1971.  Potential  of  opal  phytoliths  for  use in
   paleoecological reconstruction. _Quaternary Research_ 1: 345-
   359.
Rovner, I. 1983. Plant opal phytolith analysis:  major  advances
   in  archaeobotanical  esearch,  Pp.  225-266 in: Schiffer, M.
   (ed.) _Advances in  Archaeological  Method  and  Theory  (6)_
   Academic Press, New York.
Rovner,  I.  &  J.C.  Russ. 1992. Darwin and design in phytolith
   sytematics: Morphometric methods for  mitigating  redundancy.
   Pp.   253-276  in:  Mulholland  S.  &  G.  Rapp,  Jr.  (eds.)
   _Phytolith Systematics: Emerging issues._ Plenum  Press,  New
   York and London.
Russ,  J.C. & I. Rovner. 1987. Stereological verification of Zea
   phytolith taxonomy. _Phytolitharien Newsletter_ 4: 10-18.
Sangster, A.G. 1970. Intracellular silica deposition in immature
   leaves in three species of the Gramineae. _Annals of  Botany_
   34: 245-257.
Soni,  S.L.  & D.W. Parry. 1973. Electron probe microanalysis of
   silicon deposition in the inflorescence bracts  of  the  rice
   plant.  (_Oryza  sativa_).  _American  Journal of Botany_ 60:
   111-116.
Twiss, P.C. 1987. Grass-opal phytoliths as  climatic  indicators
   of  the  Great  Plains  Pleistocene, Pp. 179-188 in: Johnson,
   W.C.  (ed.)  _Quaternary  Environments  of  Kansas._   Kansas
   Geological Survey Guidebook Series 5. 179-188.
Twiss,  P.C., E. Suess, & R.M. Smith. 1969. Morphological class-
   ification of  grass  phytoliths.  _Soil  Science  Society  of
   America Proceedings_ 33: 109-115.


PHYTOLITH STUDIES IN WESTERN NORTH AMERICA
From: Mikhail Blinnikov [mblinnikov at stcloudstate.edu]

Virtues and values of phytolith studies

Phytoliths  are  silicified  replicas  of plant cells, which are
morphologically distinct, abundant, and durable in soils, loess,
cave deposits and other dry environments (Piperno,  1988).  Opal
phytoliths  have been described from many North American plants,
mostly  grasses  and  trees,  including  those  in  the  Pacific
Northwest  (Norgren,  1973;  Klein  and Geis, 1978; Brown, 1984;
Mulholland,  1989).  The  phytoliths'  main  strength  is  their
durability  under  a  wide  range of depositional conditions and
possibility of  identifying  plant  communities,  and  sometimes
individual  taxa, based on matching paleoassemblages with modern
analogues. In addition to  individual  shape  counts,  phytolith
concentrations can be used to infer presence/absence of forested
vegetation,  as  well  as  to indicate presence of buried A soil
horizons (Verma and Rust, 1969; Wilding and Drees,  1971).  Most
work  on phytoliths in North America has focused on archaeologi-
cal applications, it is only  recently  that  we  began  to  ap-
preciate    their   large   potential   for   paleoenvironmental
reconstructions.

Research approach (extraction, analysis, etc.)

The extraction of phytoliths from plant tissue is  done  by  dry
oxidation  for a few hours in a muffle furnace at 550 ?C, by wet
oxidation with a heated strong acid, or  combined  wet  and  dry
oxidation  (Pearsall, 2001). In my work I use modified method of
Piperno (1988, modified in Blinnikov, 1999)  for  extraction  of
phytoliths  from  soils.  Phytoliths  can  be quantitatively ex-
tracted from the silt fraction of soil or loess (5-100  microns,
20-50  g  of  soil  per  sample).  After  soil fractionation and
removal of clay and sand, the organics are  removed  by  a  con-
centrated  (70%  or  more) nitric acid with a pinch of potassium
perchlorate added per 50 ml test tube. Carbonates are removed by
mild  hydrochloric  acid.  Opal  fraction  (phytoliths)  is   be
separated from quartz and other heavier minerals using flotation
in  sodium  polytungstate  or  zinc  bromide  solution (specific
gravity of 2.3 g/cm3). Concentration of phytoliths is calculated
based on the ratio of the total dry weight of the  opal  residue
to the total dry weight of the initial sample. Information about
vegetation composition can be then obtained based on identifica-
tion  of individual phytolith shapes from fossil samples under a
light microscope and matching  the  paleoassemblages  against  a
reference  collection  from  modern  plants  and  soils by using
squared chord distance  approach  and  detrended  correspondence
analysis, or similar multivariate techniques.

Phytoliths in western North America

While  there  exists  a  considerable  bibliography on phytolith
research worldwide (Runge, 1998), little work has been  reported
from  western  North America. Some early works include Witty and
Knox (1964), Blackman (1971), Norgren (1973) and Bombin  (1984).
More recently, Blinnikov et al. (2001a, 2001b) demonstrated that
phytoliths  leave  distinct signatures under eight main types of
the forest and steppe communities of  the  Columbia  Basin,  WA.
Specifically, ponderosa pine forests can be easily distinguished
based  on the presence of diagnostic ponderosa pine "spiny body"
phytolith (see Kerns, 2001 for illustration  and  details).  Fir
and  spruce-dominated  forests  can  be  distinguished  based on
presence of silicified tracheids and  blocky  cells  of  spruce.
Potential  also  exists  in distinguishing Douglas-fir dominated
forests based on diagnostic branched asterosclereids of  Pseudo-
tsuga (Norgren, 1973).

Blinnikov  et  al.  (2001a)  also  suggests  that  phytolith as-
semblages of three types of grassland and a shrub steppe can  be
differentiated.  Because  almost  all  of  the  northwestern  US
grasses are C3 species producing mostly festucoid  rondels,  the
classical  scheme of Twiss et al. (1969) differentiating between
panicoids,  chloridoids  and  festucoids,  is  of  little   use.
However,   we   found   that   drier  _Agropyron-Poa_  dominated
grasslands have on average much smaller  percentage  of  rondels
than  more  mesic  _Festuca-Koeleria_ dominated grasslands. Fur-
thermore, any grasslands with the presence of _Stipa s.l._  will
be differentiated based on the presence of distinct _Stipa_-type
bilobate  form  which  is  different  from  "classical" panicoid
bilobates. _Aristida_ spp. can be distinguished based  on  long-
shafted  bilobate bodies also described from Aristida in Arizona
and Australia (Kerns, 2001; Bowdery, 1998).  Finer  distinctions
between,  e.g., _Koeleria_ and _Poa_, or between _Calamagrostis_
and _Bromus_ may be achieved  (Kerns,  2001;  Blinnikov,  1994).
Shrublands  with  presence of _Artemisia tridentata_ and related
sagebrush species can be separated  based  on  the  presence  of
abundant  blocky  forms  and  fragments  of  silicified  sinuous
epidermis common among  dicots,  but  not  grasses.  Communities
overrun  by  cheat  brome  (_Bromus  tectorum_) could be distin-
guished based on high percentage of silicified  epidermis  forms
is soils, common also in domesticated grasses.

Little  work  has  been  done with phytoliths from western North
American wetlands, but studies of sedge phytoliths in Russia and
the Mid-West of the US (Bobrov et al.,  2001;  Ollendorf,  1992)
suggest  that  some  sedges  can be distinguished based on their
phytoliths. Bozarth (1993) and other authors suggest that  other
phytolith  producers  among  temperate  species include nettles,
elms, oaks, sunflower family and other dicots.

Overall it appears that opal phytoliths can make the most  valu-
able  contribution  when  used  in  combination with other proxy
sources of paleoenvironmental data,  such  as  pollen,  stomata,
macrofossils,  charcoal,  and  isotope  analysis.  Due  to  con-
siderable redundancy and multiplicity  of  individual  phytolith
shapes  it  is  overall  unlikely  that  we will ever be able to
identify individual species of  plants  based  solely  on  their
phytoliths.   Differentiation   of   some   genera   of  grasses
(_Koeleria_,   _Calamagrostis_,   _Festuca_,   _Poa_,   _Stipa_,
_Aristida_)  and subgeneric level identifications in sedges are,
on the other hand, entirely possible. More productive  seems  to
be  to  search for unique community signatures in soils based on
all plants found in a particular ecosystem.

Phytoliths in western US can provide valuable information  about
the history of paleoenvironments. Some directions of future work
should include:

 1. enlarging regional phytolith collection both from individual
    species  and signatures from soils under distinct native and
    non-native plant communities
 2. fine-tuning  existing  phytolith  classifications  for   the
    region  to  include  all  major  phytolith  morphotypes from
    grasses, forbs, and trees
 3. expanding paleoenvironmental phytolith research into British
    Columbia, Alberta, and Alaska
 4. analyzing phytoliths from forested environments and wetlands
    in greater detail
 5. doing  fine  scale  phytolith  studies  to   resolve   local
    variability,  taphonomy  and  post-  depositional  transport
    issues
 6. exploring possibility of using phytoliths as a direct  proxy
    for  paleoclimates  bypassing  vegetation reconstructions by
    using transfer functions, similarly to how it is  done  with
    pollen  (e.g.,  Webb  et  al., 1998; see a Phytolith-related
    attempt in Fredlund and Tieszen, 1997),  or  stable  isotope
    analysis (Stevenson, 1997).


References

Blackman, E. 1971. Opaline silica bodies in the range grasses of
   southern Alberta. _Canadian Journal of Botany_ 49: 769-781.
Blinnikov, M. 1994. Phytolith analysis and the Holocene dynamics
   of  alpine  vegetation.  Pp.  23-40  in:  Onipchenko, V. & M.
   Blinnikov (eds.) _Experimental Investigation of Alpine  Plant
   Communities in the Northwestern Caucasus._ Veroffentlichungen
   des   Geobotanischen  Institutes  der  ETH,  Stiftung  Rbel,
   Zurich, H. 115.
Blinnikov, M., A. Busacca, & C.  Whitlock.  (2001a,  in  press).
   Reconstruction   of   the  Late  Pleistocene  Columbia  Basin
   Grassland, Washington, USA, Based  on  Phytolith  Records  in
   Loess.  _Palaeogeography,  Palaeoclimatology,  Palaeoecology_
   2714: 1-25.
Blinnikov, M., A. Busacca, & C. Whitlock. 2001b. A new  100,000-
   yr.  record from the Columbia Basin, Washington, USA. Pp. 27-
   55 in: J. D. Meunier, J.D. &  F.  Colin  (eds.)  _Phytoliths:
   Applications  in  earth  sciences  and  human history._ A. A.
   Balkema, Rotterdam, the Netherlands.
Bobrov, A. A., Bobrova, E. K., Alexeev  Ju.  E.  2001.  Biogenic
   silica  in  biosystematics:  potential  uses. Pp. 279-288 in:
   Meunier J.D. & F. Colin (eds.) _Phytoliths:  Applications  in
   Earth  Sciences and Human History._ A. A. Balkema, Rotterdam,
   the Netherlands.
Bowdery, D. 1998. _Phytolith Analysis  Applied  to  Pleistocene-
   Holocene  Archaeological  Sites in the Australian Arid Zone._
   British Archaeological Reports International Series, v. 695.
Bozarth, S. R. 1993. Biosilicate assemblages of  boreal  forests
   and  aspen  parklands.  Pp.  95-105 in: Pearsall, D.M. & D.R.
   Piperno  (eds.)  _Current  Research  in  Phytolith  Analysis:
   Applications  in Archaeology and Paleoecology_ MASCA Research
   Papers in Science and Archaeology, Volume 10.
Brown, D. 1984. Prospects and limits  of  a  Phytolith  key  for
   grasses  in  the  central  United  States.  _Journal  of  Ar-
   chaeological Sciences_ 11: 345-368.
Bombin, M. 1984. _On  phytoliths,  late  Quaternary  ecology  of
   Beringia,  and  information evolutionary theory._ Unpublished
   Ph.D. dissertation, University of Alberta, Calgary,  Alberta,
   Canada. 163 p.
Fredlund,  G.  G.,  &  L.L.  Tieszen.  1997.  Calibrating  grass
   phytolith assemblages in climatic terms: Application to  late
   Pleistocene    assemblages    from   Kansas   and   Nebraska.
   _Palaeogeography, Palaeoclimatology, Palaeoecology_ 136: 199-
   211.
Kearns, B. 2001. Diagnostic phytoliths  for  a  ponderosa  pine-
   bunchgrass  community near Flagstaff, Arizona. _The Southwes-
   tern Naturalist_ 46(2): 282-294.
Klein, R.  L.,  &  J.W.  Geis.  1978.  Biogenetic  opal  in  the
   Pinaceae. _Soil Science_ 126: 145-156.
Mulholland,  S.  C.  1989.  Phytolith shape frequencies in North
   Dakota grasses: a comparison to general patterns. _Journal of
   Archaeological Science_ 16: 489-511.
Norgren, J. A. 1973. _Distribution,  Form  and  Significance  of
   Plant Opal in Oregon Soils._ Ph.D. dissertation, Oregon State
   University, Corvallis, OR.
Ollendorf,  A.  L. 1992. Toward a classification scheme of sedge
   (Cyperaceae) phytoliths. Pp. 91-106 in: Mulholland  S.  &  G.
   Rapp,  Jr.  (eds.)  _Phytolith Systematics: Emerging issues._
   Plenum Press, New York, NY.
Pearsall, D. 2001. _Paleoethnobotany: A Handbook of Procedrues._
   2nd ed. Academic Press, San Diego, CA.
Piperno, D. 1988.  _Phytolith  Analysis:  An  Archeological  and
   Geological Perspective._ Academic Press, San Diego, CA.
Runge, F. 1998. _Bibliography of Phytolith Research._ University
   of Paderborn, Germany.
Stevenson,  B.  A.  1997.  _Stable Carbon and Oxygen Isotopes in
   Soils and Paleosols of the Palouse Loess, Eastern  Washington
   State:    Modern    Relationships    and   Applications   for
   Paleoclimatic Reconstruction._  Unpublished  Ph.D.  disserta-
   tion, Colorado State University, Fort Collins, CO.
Twiss,  P.C.,  Suess,C.E.,  and  Smith, R.M. 1969. Morphological
   classification of grass phytoliths. _Soil Science Society  of
   America Proceedings_ 33: 109-115.
Verma,  S. D. & R. H. Rust. 1969. Observation on opal phytoliths
   in a soil biosequence in southeastern Minnesota.  _Soil  Sci.
   Soc. America Proc._ 33: 749-751.
Webb,  T.  III, K.H. Anderson, P.J. Bartlein, & R.S. Webb. 1998.
   Late Quaternary climate change in eastern  North  America:  a
   comparison  of  pollen-derived  estimates  with climate model
   results. _Quaternary Science Reviews_ 17: 587-606.
Wilding, L. P. & L.R. Drees. 1971. Biogenic opal in Ohio  soils.
   _Soil Sci. Soc. America Proc._ 35: 1004-1010.
Witty,  J.  E.  &  E.G.  Knox,  E.  G.  1964. Grass opal in some
   Chestnut and Forest soils  in  north  central  Oregon.  _Soil
   Science Society of America Proceedings_ 28: 685-687.

Phytoliths on line:

Mikhail Blinnikov's phytolith gallery:
http://coss.stcloudstate.edu/mblinnikov/phd/phyt.html

Terry Ball's phytolith page:
http://reled.byu.edu/ascript/tball/index2.html
[this link did not work on 2002/02/20]

Deb Pearsal's phytolith web site:
http://web.missouri.edu/~phyto/

University of Arizona phytolith links:
http://www.geo.arizona.edu/palynology/pphyolth.html

Glen Fredlund's webpage:
http://www.uwm.edu/People//fredlund/phytocat.htm


ULTIMATE PALINDROME
From: "Scott D. Russell" [srussell at ou.edu]

Dear Adolf:

Here's  a  true  challenge.  I think you should rush to press at
8:02 pm tonight! Here is an interesting palindrome as the excuse
(which I got from my stepbrother who got from someone else  ....
etc!):

Believe  it  or  not, 8.02pm on February 20 this year will be an
historic moment in time. It will not be marked by the chiming of
any clocks or theringing of bells, but at that precise time,  on
that specific date, something will happen which has not occurred
for  1,001 years and will never happen again. As the clock ticks
over from 8.01pm on Wednesday, February 20, time will, for sixty
seconds only, read in perfect symmetry 2002, 2002, 2002,  or  to
be  more  precise  -  20:02, 20/02, 2002. The last occasion that
time read in such a symmetrical pattern was long before the days
of the digital watch and the 24-hour clock at 10.01am on January
10, 1001. And because the clock only goes up  to  23.59,  it  is
something that will NEVER happen again.


Sorry, Scott!
At  this  palindromic  moment  we  will  be [again] in Portland,
Oregon, and I will be sipping Pilsener Urquell with Oluna and my
friends in the Rheinlander. - Adolf
P.S. When I looked at the date posted in BEN, I realized that  I
have  not  changed  the  year in the two previous BEN issues and
they are labelled as "2001". Can  you change it in the web page? 
Thanks. - AC

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