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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
LIFE, DEATH AND ORIGINS OF ANCIENT ICEMEN: PALAEOFORENSIC BOTANY
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
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,
[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.
COSTS AND CONSEQUENCES OF NON-INDIGENOUS PLANTS
IN THE BRITISH ISLES
From: Mark Williamson, Department of Biology,
University of York, York YO10 5DD, UK
[mw1 at york.ac.uk]
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.
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 &
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_,
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
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
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).
_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
BOTANIST JOB POSTING: ROYAL BOTANICAL GARDENS HAMILTON, ONTARIO
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
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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