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BEN # 281

Adolf Ceska aceska at victoria.tc.ca
Fri Feb 1 02:35:06 EST 2002

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No. 281                              January 31, 2001

aceska at victoria.tc.ca                Victoria, B.C.
 Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2

From: James Holms Dickson [J.Dickson at bio.gla.ac.uk]

By  "iceman"  I  mean  an  ancient  male  human body melted from
glacier ice, as in the case  of  the  5,300  year  old  Tyrolean
Iceman, or Oetzi as he was quickly nicknamed after the discovery
high  in  the Alps in 1991. Oetzi's corpse has been studied by a
great variety of scientific techniques  among  which  investiga-
tions  of  both microscopic and macroscopic plant remains, found
both internally and externally,  have  revealed  much  on  diet,
other  aspects  of  ethnobotany, season of death and environment
(contributions by Dickson, Oeggl, and Rott in  Bortenschlager  &
Oeggl  2000;  Dickson 2000; Fowler 2000). We know what Oetzi ate
during the last few days of his life, we think that his homeland
was to the south not to the north, we think his  provisions  may
have  been  wrapped  in  moss,  we think he died in Spring/early
Summer not in autumn as first  claimed  and  these  studies  and
others are continuing and much has changed recently.

The  first  ever North American Iceman is Long Ago Person Found,
Kwaday Dan Ts'nchi in the local First Nation language  (KDT  for
short).  He  was a male of about twenty years when he died about
550 years ago. His remains were found on the  14th  August  1999
protruding  from  a  melting, lateral part of the Samuel Glacier
(Beattie et al. 2000). [At the very moment of  discovery  I  was
about 50 miles away!]

The  elevation  of  the site is approximately 1500 m (about 5000
feet) and the place is Fault  Creek  in  the  Tatshenshini-Alsek
Provincial  Park in northernmost British Columbia (59 deg. N 136
deg. W). KDT had lived and died long before Europeans penetrated
into the region and he died on a full stomach.

Such icemen as Oetzi and KDT pose questions different from those
arising from frozen burials such as  those  of  the  Inuit,  the
Siberians  in tombs, the sacrificed Andean children, or even the
lead-poisoned sailors of the Franklin expedition. What  had  the
icemen  been  doing where they were found, had they traveled far
or not, what exactly were the events  immediately  before  death
and  what  happened  thereafter,  what  were  their  racial  and
geographic affinities? In the case of  Oetzi  we  know  some  of
these  answers and for KDT we are in the process of finding out.
Richard Hebda of the Royal British Columbia  Museum  and  I  are
working away merrily on the plant remains (pollen, seeds, leaves
and mosses) and interesting findings are emerging.


Beattie,  O.,  B. Apland, E. Blake, J.A. Coswgrove, S. Gaunt, S.
   Greer, A.P. Mackie, K.E. Mackie, D. Straathof,  V.  Thorp,  &
   P.M.  Troffe.  2000.  The Kwaday Dan Ts'nchi discovery from a
   glacier in British Columbia. Canadian Journal of  Archaeology
   24, 129-147.
Bortenschlager,  S.  & K. Oeggl. (eds.) 2000. _The Iceman in his
   natural  environment:  palaeobotanical   results._   Springer
   Verlag, Wien & New York.
Dickson,  J.H., K. Oeggl, T. Holden, L. Handley, T.C. O'Connell,
   & T. Preston. 2000. The  omnivorous  Tyrolean  Iceman:  colon
   contents  (cereals,  meat,  moss  and  whipworm)  and  stable
   isotopes. _Philosophical Transactions of the Royal Society of
   London B_ 355: 1-7.
Fowler, Barbara. 2000. _Iceman: uncovering the life and times of
   a prehistoric man found in an alpine glacier._ Random  House,
   New York.
   [This  is  a  racy  account by a journalist who thinks that I
   work in " The University of Scotland"!]

Dr. Jim Dickson,  Professor  of  Archaeobotany  and  Plant  Sys-
tematics in the University of Glasgow, will talk about "Life and
death  of  the  Tyrolean  Iceman: clues from plants" in Victoria
B.C., on Tuesday, February 19, 2002, 7:30 p.m., UVIC  -  Elliott
168. This lecture is a part of the Botany Night series organized
by the Victoria Natural History Society. Admission is free.

From: Mark Williamson, Department of Biology,
   University of York, York YO10 5DD, UK
   [mw1 at york.ac.uk]

The problem

Alien  species  are a world-wide problem of much concern. Nobody
can doubt the supreme importance of controlling aliens in,  say,
Hawaii  or  New  Zealand. Many popular books give the impression
that all introduced species eventually become problems. However,
the impact of introduced species does vary from place  to  place
and  from  group to group. Here I want to consider just vascular
plants in the British Isles.

The numbers

It is customary to distinguish different stages of  invasion.  A
plant  may be brought into a country alive but not found outside
cultivation. It could just occur as seeds  in  a  herbarium,  it
could  be  grown in a botanic or other garden. This stage I have
called 'imported' (Williamson 1996). The next stage is for it to
escape captivity, often called, for plants, 'casual' (Richardson
et al. 2000). A short or long time later, it may be  found  that
the  plant  species  has  a  fully  established, self-sustaining
population, the 'established' stage.  Some  established  species
are,  for various reasons, disliked, attacked, controlled. These
are usually called weeds, or pests. Weeds  may  also  be  casual
species  as  in  volunteers  from some crops. It is common (e.g.
Lonsdale 1999) to compare the numbers of non-indigenous or alien
species with the numbers of natives,  often,  but  by  no  means
always, using the number of established aliens.

The British flora is as well known as any. What are the numbers?

Excluding  micro-species,  the number of natives is around 1400.
There are 1350 listed in the forthcoming Atlas 2000 (Preston  et
al. in press), 1407 in the Ecological Flora Database (Fitter and
Peat  1994),  1481  in  Preston  & Hill (1997). Other counts are
given in Williamson (in press). One reason for the variation  is
uncertainty  about  which  are native and which introduced, par-
ticularly which were introduced long  ago.  A  central  European
habit  is  to talk of archaeophytes for plants introduced before
1492  or  thereabouts  (which  excludes  American  species)  and
neophytes  (a  word  with  other  meanings) for those introduced
later. The Atlas 2000 is the first time that  this  concept  has
been  used  in  a  book  in  Britain and some species previously
listed as native have been reclassified.  This  has  caused  un-
necessary  alarm  in  some  conservation  circles, partly from a
misunderstanding  that  the  Convention  on  Biodiversity  urges
countries to control all aliens. What Article 8 actually says is
"as far as possible and as appropriate ... prevent the introduc-
tion of, control or eradicate those alien species which threaten
ecosystems,  habitats or species" which refers, as will be seen,
to a small sub-set of  aliens  and  even  so  allows  plenty  of

Counts  of  British  aliens are much more variable than those of
natives, partly because different botanists have very  different
ideas  of  where to draw the line between casual and established
and of whether casuals should be included. In Williamson  (1993)
I  distinguished  between widely naturalized (67 species), fully
naturalized(143  more)  and  locally  naturalized  (348   more),
naturalized  being  a  term often equivalent, as here, to estab-
lished,  but  often  meaning  growing   in   native   vegetation
(Richardson et al. 2000). The fully naturalised figure of 210 is
quite close to the number in the Ecological Flora Database (196)
(Fitter and Peat 1994) and in the British Ellenberg tables (239)
(Hill  at al. 1999). My locally-or-more naturalized number (558)
is close to the numbers in the records  behind  the  1958  Atlas
(Perring  &  Walters  1962) and the 1988 Sample Survey (Palmer &
Bratton 1995), namely 588 in 1958, 605 in 1988, 468 in both, 725
in either (Williamson et al. in press). But we  found  only  118
neophytes  reliably  mapped in both those surveys (Williamson et
al. in press). If casuals are included  the  total  numbers  are
much larger, 1642 in Williamson (1993), 3467 in Clement & Foster
(1994) with Ryves et al. (1996).

For  the Atlas 2000 project (Preston et. al. in press) there are
numbers for the species mapped in the book  and  listed  on  the
accompanying  CD. These are, in both, 1350 natives (as mentioned
above), 44 species which may be native or  alien,  and  147  ar-
chaeophytes. The book maps 505 neophytes, the CD lists 651 more.
Surprisingly,  the  book maps a further 40 as casuals and the CD
adds 199 more, even though many botanists would probably  regard
at least the 651 extra neophytes as casuals.

All  that  seems to me to indicate that there are in Britain (or
the British Isles, it makes  no  difference  at  this  accuracy)
about  1400  native  species,  around 200 definitely established
aliens, 300-500 more that may be established somewhere, and some
thousands more that have been recorded as casuals at some time.

There are two further complications, one to the totals,  one  to
the  definitions.  To  the  totals  could be added microspecies,
mostly in three genera.  Stace  (1997)  states  that  there  are
currently  261  such  listed  in  _Hieracium_, including "a con-
siderable number of aliens", 320 in _Rubus_,  apparently  almost
all  native,  and 229 in _Taraxacum_, probably over half of them
alien. At a guess that adds about 600  natives  and  200  estab-
lished aliens, quite a big addition.

The  other  complication comes from the distinction between non-
indigenous and alien. _Spartina anglica_ is  the  allotetraploid
formed  in  England  late  in  the 19th century from _Spartina x
townesendii_, the hybrid, formed earlier in  the  19th  century,
between  the  native  _S.  maritima_  and the American alien _S.
alterniflora_. British floras list _Spartina anglica_ as native.
It is not alien (brought in  from  elsewhere)  but  it  is  non-
indigenous  (not found in Britain in the Mesolithic period). The
German term is heimatlos (homeless), with no native range on one
obvious interpretation. Two notable  species  listed  as  aliens
should  perhaps  be  listed in the same way. _Senecio squalidus_
(Oxford ragwort) arose in the Oxford Botanic Gardens in the 17th
and 18th centuries from hybrids collected in Sicily  (Abbott  et
al.  2000).  British _Rhododendron ponticum_ is at least partly,
possibly largely, hybrid, formed in Britain between Spanish  _R.
ponticum_,  American _R. catawbiense_ and other species (Milne &
Abbott 2000).

Cost and benefit estimates

The concern about aliens is, broadly, about their  environmental
and  economic  costs.  Some,  notably  some crop plants, produce
considerable economic benefits. Any benefits should be  balanced
against costs. Valuing environmental effects is well known to be
difficult (Perrings et al. 2000).

On  the cost side, there has been one useful attempt to estimate
the control costs of all British plants treated fully in  floras
(Prus  1996). From the national costs of herbicides, information
on which plants were controlled by which herbicide, supplemented
by information on other forms of control, he was able to  derive
a  figure,  in  pounds  sterling  per  year, for effectively all
species. A graphical presentation of his results  for  the  more
costly  species  classified as native, archaeophyte and neophyte
is given in Williamson (1998). More recently  I  (Williamson  in
press)  have  tried to estimate the costs and benefits of thirty
interesting (widespread, potential pests,  etc.)  non-indigenous
species.  The  Prus cost for these varies from 58 million pounds
to a fraction of a pound. Clearly the cheaper  ones  are  in  no
sense  a  pest on this measure, indeed only nine score more than
GBP 22,000, which is still a trivial cost nationally.  Indeed  I
cannot  take  two  of  them (_Galinsoga parviflora_, _Matricaria
discoidea_) seriously as threatening invasives in the CBD sense,
but   two   others   (_Heracleum   mantegazzianum_,   _Impatiens
glandulifera_)  represent  more serious threats not reflected in
their Prus cost. That makes just nine alien species which are in
some sense threatening in the British Isles.

These nine species can be classified as:

 1. three annual arable weeds  (_Avena  fatua_,  _A.  sterilis_,
    _Veronica persica_),
 2. one   serious  garden  weed,  an  archaeophyte  (_Aegopodium
 3. three weeds of forest  and  woodland  (and  more)  but  with
    compensating  benefits  (_Acer  pseudoplatanus_,  _Impatiens
    glandulifera_, _Rhododendron ponticum_), and
 4. two  pests  of  other  habitats  and  officially  proscribed
    (_Fallopia japonica_, _Heracleum mantegazzianum_).

That's  a  remarkably  small  proportion of however many hundred
species are regarded as established  non-indigenous  species.  A
few tens more might sometimes be regarded as weeds (like _Galin-
soga  parviflora_  and  _Matricaria discoidea_ mentioned above).
Williamson & Brown (1986) counted  39.  Depending  on  what  you
regard  as  a  pest  and  what as established, the proportion of
current pests among the currently established seems to  be  only
between one and ten percent.

Other impacts and time lags

The  cost  of control is far from being the only impact that can
be  measured.  In  Williamson  (1998)  I  reported  five   other
measurable  impacts.  Briefly,  British established alien plants
have, species for species, the same impact as native plants when
measured as weediness in various ways or as local abundance  but
have  a  lesser impact measured as range. This is partly because
most British aliens are still increasing their range.  Typically
it  takes  a  century or two to achieve a full range in Britain;
curves for the spread of three alien _Impatiens_  are  given  in
Williamson (1996) which demonstrate this.

Of  the  thirty interesting non-indigenous species considered in
Williamson (in press) - see Appendix below - all  except  eight,
i.e. 22, have significant increases from 1958 to 1988. The eight
are  _Acer pseudoplatanus_, _Aegopodium podagraria_, _Cymbalaria
muralis_,  _Galinsoga   parviflora_,   _Matricaria   discoidea_,
_Mimulus   guttatus  s.l._,  _Spartina  anglica_  and  _Veronica
persica_. These are  all  introductions  of  the  mid-nineteenth
century or earlier except _Spartina anglica_ which has been much
spread  deliberately.  All  eight  are widespread in the British
Isles except _Galinsoga  parviflora_  which  is  only  found  in
central  and southern England. So some of those 22 species still
spreading may yet have much more serious impact than  they  have
had  so  far;  _Crassula  helmsii_  being a particular cause for
concern in taking over ponds and similar water  bodies.  But  as
new  species are still becoming established, it may well be that
the proportion of pests will scarcely change.


Some non-indigenous plants in Britain cause serious economic and
environmental damage. These species appear to be only one to ten
percent of  those  that  establish  permanent  populations.  The
numbers  of both established and pest species can be expected to
increase, though fairly slowly, for the foreseeable future.


Abbott, R.J., J.K. James, J.A. Irwin, & H.P. Comes. 2000. Hybrid
   origin of the Oxford Ragwort, _Senecio  squalidus_  L.  _Wat-
   sonia_ 23: 123-138.
Clement,  E.J. & M.C. Foster. 1994. _Alien Plants of the British
   Isles. A Provisional Catalogue of Vascular Plants  (excluding
   Grasses)_. Botanical Society of the British Isles, London.
Fitter,  A.H.  & H.J. Peat. 1994. The ecological flora database.
   _Journal of Ecology_  82: 415-425.
Hill,  M.O.,  J.O.  Mountford,  D.B.  Roy, & R.G.H. Bunce. 1999.
   Ellenberg's Indicator Values for British Plants. Institute of
   Terrestrial Ecology, Huntingdon.
Lonsdale, W.M. 1999. Global patterns of plant invasions and  the
   concept of invasibility. _Ecology_ 80: 1522-1536.
Milne,  R.I.  &  R.J.  Abbott. 2000. Origin and evolution of in-
   vasive naturalized material of _Rhododendron ponticum_ L.  in
   the British Isles. _Molecular Ecology_ 9: 541-556.
Palmer,  M. A. and Bratton J. H. (eds.) 1995. A Sample Survey of
   the Flora of Britain and Ireland: The  Botanical  Society  of
   the  British  Isles  Monitoring  Scheme  1987-1988. UK Nature
   Conservation No.  8,  Joint  Nature  Conservation  Committee,
Perring,  F.H.  &  S.M.  Walters.  1962.  _Atlas  of the British
   Flora._ Thomas Nelson and Sons, London.
Perrings, C., M. Williamson, & S. Dalmazzone. (eds.) 2000.  _The
   Economics of Biological Invasions._ Edward Elgar, Cheltenham.
Preston,  C.D. & M.O. Hill. 1997. The geographical relationships
   of British and Irish vascular plants. _Botanical  Journal  of
   the Linnean Society_ 124: 1-120.
Preston, C.D., D.A. Pearman, & T.D. Dines. (eds.) in press. _New
   Atlas  of  the  British  and  Irish Flora._ Oxford University
   Press, Oxford.
Prus, J.L. 1996. _New methods of risk assessment for the release
   of transgenic plants._ PhD thesis, Cranfield University.
Richardson, D.M., P. Pysek,  M.  Rejmanek,  M.G.  Barbour,  F.D.
   Panetta,  &  C.J.  West. 2000. Naturalization and invasion of
   alien plants: concepts and definitions. _Diversity  and  Dis-
   tributions_ 6: 93-107.
Ryves,  T.B.,  E.J. Clement, & M.C. Foster. 1996. _Alien Grasses
   of the British  Isles._  Botanical  Society  of  the  British
   Isles, London.
Stace,  C.  1997. _New Flora of the British Isles._ 2nd edition.
   Cambridge University Press, Cambridge.
Williamson, M. 1993. Invaders, weeds and the risk  from  geneti-
   cally manipulated organisms. _Experientia_ 49, 219-224.
Williamson,  M.  1996.  _Biological  Invasions._ Chapman & Hall,
Williamson, M. 1998. Measuring the impact of plant  invaders  in
   Britain. Pp. 57-68 in Starfinger, U., K. Edwards, I. Kowarik,
   &   M.   Williamson.   (eds).  _Plant  Invasions:  Ecological
   Mechanisms and Human Responses._ Backhuys, Leiden.
Williamson, M. in press. Alien plants in the British Isles.  In:
   Pimentel, D. (ed.) _Environmental and Economic Costs of Alien
   Plant,  Animal and Microbe Invasions_. CRC Press, Boca Raton,
Williamson, M. & K.C. Brown. 1986. The analysis and modelling of
   British invasions. _Philosophical Transactions of  the  Royal
   Society_ B 314: 505-522.
Williamson,  M., C. Preston, & M. Telfer. in press. On the rates
   of spread of British alien plants. In Child, L. et al. (eds.)
   _Plant Invasions: Proceedings of the 6th  International  Con-
   ference on Invasive Alien Plants._ Backhuys, Leiden.


The  thirty interesting non-indigenous British plants considered
in Williamson (in press).

Species Family

_Acer pseudoplatanus_ Aceraceae
_Aegopodium podagraria_ Apiaceae
_Allium triquetrum_ Liliaceae
_Avena fatua_ Poaceae
_Avena sterilis_ Poaceae

_Buddleja davidii_ Buddlejaceae
_Centranthus ruber_ Valerianaceae
_Conyza canadensis_ Asteraceae
_Crassula helmsii_ Crassulaceae
_Crepis vesicaria_ Asteraceae

_Cymbalaria muralis_ Scrophulariaceae
_Elodea canadensis_ Hydrocharitaceae
_Elodea nuttallii_ Hydrocharitaceae
_Epilobium brunnescens_ Onagraceae
_Epilobium ciliatum_ Onagraceae

_Fallopia japonica_ Polygonaceae
_Galinsoga parviflora_ Asteraceae
_Gunnera tinctoria_ Gunneraceae
_Heracleum mantegazzianum_ Apiaceae
_Impatiens glandulifera_ Balsaminaceae

_Matricaria discoidea_ Asteraceae
_Mimulus guttatus_ Scrophulariaceae
_Quercus ilex_ Fagaceae
_Rhododendron ponticum_ Ericaceae
_Senecio squalidus_ Asteraceae

_Smyrnium olusatrum_ Apiaceae
_Spartina anglica_ Poaceaea
_Symphoricarpos albus_ Caprifoliaceae
_Veronica filiformis_ Scrophulariaceae
_Veronica persica_ Scrophulariaceae

From: David Galbraith [galbraith at rbg.ca]

Royal Botanical Gardens, Canada's largest botanical garden whose
holdings include 1100 ha of formal gardens, wetlands  and  wood-
lands is seeking a Field Botanist. This a two year term position
[the incumbent is on a leave of absence] and begins on April 1st
2001.  The  successful  candidate will have at least 5 years ex-
perience in field botany, herbarium curation and  documentation,
supervision of students and interns and managing a budget.

The  ideal  candidate  will also be a team player who is innova-
tive, creative and finds work  challenging  and  rewarding.  Ex-
perience in preparing budgets is also an asset. An undergraduate
degree in botany is essential.

Please  forward  your resume in confidence by February 18th 2002

   Susan Ingram
   Director, Human resources
   Royal Botanical Gardens
   P.O. Box 399
   Hamilton, Ontario L8N 3H8
   Fax 905 577- 0375
   E-mail: hire at rbg.ca

We thank all applicants: however, only  those  selected  for  an
interview will be contacted. [Not very nice, eh? - AC]

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