BEN # 209 - Dr. W.A. Weber Festschrift - Part III

Adolf Ceska aceska at VICTORIA.TC.CA
Tue Nov 24 04:23:43 EST 1998


                                                   
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No. 209                              November 24, 1998

aceska at victoria.tc.ca                Victoria, B.C.
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 Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2
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            BEN # 207, 208, and 209 are dedicated to
                  the doyen of Colorado botany

                DR. WILLIAM ALFRED (BILL) WEBER

    on the occasion of his 80th birthday, November 16, 1998.


GENUS CRATAEGUS (HAWTHORN) FROM ALASKA TO CALIFORNIA
From: J.B. Phipps <jphipps at julian.uwo.ca>

This  paper  covers the known Crataegus L. (Rosaceae) species of
Alaska, British Columbia, Oregon, Washington, California,  Idaho
and  Montana. Recent discoveries based on extensive fieldwork by
the author and R.J. O'Kennon in the western states  and  British
Columbia,  insights  coming  from  the  biosystematic studies of
Dickinson and associates and work by Brunsfeld  and  Johnson  on
C.douglasii  and  relatives  have radically altered the views of
Crataegus of this area previously held to  comprise  only  three
native species: one black-fruited - C. douglasii Lindl. with its
two  varieties,  var.  douglasii  and  var.  suksdorfii  (Sarg.)
Kruschke; and two red-fruited - the C. piperi  Britton  of  this
region,  but  generally called C. columbiana, and C. macracantha
Lodd. ex Loud. reported for Montana, but  frequently  called  C.
succulenta  Schrad.  ex  Link  or C. occidentalis Britton in the
west. In addition the red-fruited introduction C. monogyna Jacq.
was widely reported. My work (Phipps, 1998; Phipps and O'Kennon,
1998) greatly extends the range of C. macracantha and adds three
more species for the region. I believe  that  further  novelties
are  still  to  be  expected  here.  I  give  below a key to the
Crataegus taxa, shortness of notice  for  preparation  of  which
(only  one  week)  has precluded detailed testing, together with
brief notes on some of the species. It is  interesting  to  note
that  the majority of species are in the purple to black-fruited
group.

KEY
Note: 'Leaves' refers to short-shoot leaves only.

1. Leaves with veins to the sinuses; thorns, at  least  some  of
   them,  indeterminate  and  growing out into twigs; introduced
   species:

   2. Leaves deeply 5-7 lobed, leaf incision index > 50%,  lobes
      acute; style and nutlet 1
      ..........................................  1. C. MONOGYNA

   2. Leaves  shallowly  3-lobed,  leaf  incision  index usually
      < 30%, lobes obtuse, terminal usually largest; styles  and
      nutlets 2
      .........................................  2. C. LAEVIGATA

1. Leaves  lacking  veins  to  sinuses;  thorns all determinate;
   native species:

   3. Fruit bright red at full maturity, generally more or  less
      orange a month before:

      4. Autumnal color of leaves generally yellow in this area;
         one-year  old  twigs  fawn,  golden-green  or pale tan;
         early flowering; nutlets not pitted laterally
         ...................................  10. C. CHRYSOCARPA

      4. Autumnal color of leaves bronze in this area;  one-year
         old  twigs  usually  dark purple-brown; late flowering;
         nutlets pitted laterally
         ...................................  11. C. MACRACANTHA

   3. Fruit purple to black at full maturity, in some bright red
      about a month prior to this:

      5. Leaves ovate, lobes subacute to  obtuse;  inflorescence
         branches densely hairy; fruit orbicular, hairy;
         .......................................  9. C. PHIPPSII

      5. Without above combination of characteristics:

         6. Thorns fine, usually straight; older twigs and young
            branches usually shining copper-colored; leaf blades
            generally at least 2 x as long as wide; stamens 10
            ...................................  3. C. RIVULARIS

         6. Thorns  stouter, often slightly recurved; 2-year old
            twigs usually very deep brown,  often  purple-brown;
            leaf  blades  usually  about 1.5 x as long as broad;
            stamens 10 or 20:

            7. Thorns  2-6 cm  long;  calyx   lobes   glandular-
               serrate,  acute,  erect  to  spreading  in fruit;
               fruit bright red a month  before  full  maturity,
               flask shaped:

               8. Leaves thin, sharply lobed; anthers pink; very
                  late anthesis; calyx lobes in fruit reflexed
                  ............................  7. C. WILLIAMSII

               8. Leaves coriaceous, less sharply lobed; anthers
                  white;  early  to  mid-season  anthesis; calyx
                  lobes in fruit spreading
                  .........................  8. C. OKANAGANENSIS

            7. Thorns usually about  2.5 cm  long;  calyx  lobes
               scarcely   glandular   and  not  serrate,  blunt,
               shorter, in fruit appressed;  fruit  often  black
               but occasionally vinous or chestnut-coloured (but
               not bright red) a month before full maturity:

               9. Stamens 20;   leaf  blades  obscurely  or  not
                  lobed, lobes usually blunt
                  ............................  4. C. SUKSDORFII

               9. Stamens 10;  leaf  blades   usually   clearly,
                  though  not  necessarily  very  deeply, lobed,
                  lobes usually subacute to acute:

                  10. Flowers 12-15 mm in diameter; fruit ellip-
                      soid, usually black or dark vinous a month
                      before full maturity
                      .........................  5. C. DOUGLASII

                  10. Flowers 15-18 mm in diameter; fruit ampul-
                      liform,  usually  vinous  or chestnut at a
                      month before full maturity
                      .........................  6. C. OKENNONII


NOTES ON TAXA KEYED
(extraterritorial distributions not indicated)

1. C. MONOGYNA Jacq.
   Common at low  altitudes  in  northwestern  WA,  southwestern
   coastal  BC;  scattered  elsewhere; AK, interior BC, interior
   WA, CA, OR, MT. Hybridizes locally with  C.  douglasii  (Love
   and Feigen, 1978)

2. C. LAEVIGATA (Poir.) DC.
   For  first  accurate  wild  record  of this species for North
   America see Phipps (1998). Very rare,  WA.  The  double  red-
   flowered  cultivated hawthorns usually attributed here are C.
   x media Bechst.

3. C. RIVULARIS Nutt. ap. Torr. & A. Gray
   Southern ID (fairly common)  and  southwards.  Apparently  no
   intermediates  with  C.  douglasii,  with  which range is al-
   lopatric. Quite different from latter  species  (see  Phipps,
   Sida, submitted).

4. C. SUKSDORFII (Sarg.) Kruschke
   Resuscitated  as  a  species by Brunsfeld and Johnston (1990)
   with good arguments. Rather variable, however.  AK,  BC,  WA,
   OR, CA, ID, MT.

5. C. DOUGLASII Lindl.
   Common  and  widespread  in  area  OR, WA, BC, ID, MT. Rather
   variable and segregates or varieties might be recognized.

6. C. OKENNONII J.B. Phipps
   Segregate from C. douglasii with  larger  flowers,  different
   leaves,  fruit and growth habit. For more detail see Phipps &
   O'Kennon (1998). WA, BC, ID, MT.

7. C. WILLIAMSII Eggl.
   A perfectly good species  of  northwest  MT  ignored  by  all
   floras since its description. Very late flowering and easy to
   recognize. See Phipps (1998).

8. C. OKANAGANENSIS J.B. Phipps & O'Kennon
   Surprisingly, for a new species, very common from Okanagan of
   BC  and  WA  to  northwest MT. Could only be confused with C.
   williamsii of species listed here. See  Phipps  and  O'Kennon
   (1998).  St.  John (1963) may have alluded to this species in
   his Flora of southeastern Washington and adjacent Idaho. Fred
   Johnson was the only person to collect this species  in  sub-
   stantial numbers before ourselves.

9. C. PHIPPSII O'Kennon
   The  most  distinct  of  the  species  recently described and
   scattered from BC and WA  to  MT.  See  Phipps  and  O'Kennon
   (1998).

10. C. CHRYSOCARPA Ashe
   Includes  C.  piperi  Britton  and  "C. columbiana Howell" of
   floras. See Phipps (1998) for discussion of correct  name.  I
   recognize two varieties. BC, WA, OR, ID, MT.

11. C. MACRACANTHA Lodd. ex Loud.
   O'Kennon  and  I  extend its range westward to ID, WA, OR and
   BC. It is quite common in the  Okanagan  and  easily  distin-
   guished,  at  all  seasons,  from any other Crataegus treated
   here.

I would like to take this opportunity to take my hat off  to  an
old  friend,  Bill Weber, greatly deserving of this Festschrift.
It is perhaps symbolic that Colorado, of all the western  states
(west  of  the  Rockies)  has  been  the  only  one with several
Crataegus species not to have its list altered in the  slightest
by recent work.

References:
Brunfeld,  S.J. and F.D. Johnson. 1990. Cytological, morphologi-
   cal and phenological support for specific status of Crataegus
   suksdorfii (Rosaceae). Madrono 37: 274-282.
Love, R. & M. Feigen. 1978. Interspecific hybridization  between
   native   and  naturalized  Crataegus  (Rosaceae)  in  western
   Oregon. Madrono 25: 211-217.
Phipps, J.B. 1998. Introduction to the red-fruited hawthorns  of
   western  North  America.  Canad.  J.  Bot. 76 (July issue, in
   press )
Phipps, J.B.  &  R.J.  O'Kennon.  1998.  Three  new  species  of
   Crataegus  (Rosaceae)  from  western  North America. C. oken-
   nonii, C. okanaganensis and C. phippsii. Sida 19: 169-191.
St. John, H. 1963. Flora of southeastern Washington and adjacent
   Idaho, 3rd ed. Outdoor Pictures, Escondido, California.


DISTRIBUTION OF THE CLADONIACEAE IN WESTERN NORTH AMERICA
From: Trevor Goward <tgoward at mail.wellsgray.net> and
      Teuvo Ahti <ahti at cc.helsinki.fi>

In our recent study (Goward & Ahti, 1997), we have examined  the
western  North  American distributions of 84 taxa and chemotypes
of Cladinae and Cladoniae  occurring  at  temperate  and  boreal
latitudes.  Our  analysis  was drawn primarily from the maps and
text of Goward, Ahti & Brodo (in prep.: based  on  approximately
8,200  specimens!),  and secondarily from those of Geiser et al.
(1994), Hammer (1995), and Thomson (1984). We propose six  broad
conclusions,  some  of  which may be of general interest to stu-
dents of phytogeography:

 1. Western North America's richest assemblage  of  Cladina  and
    Cladonia,  with  between  76  and 78 taxa, occurs in British
    Columbia between 52N and 56N, in a region covered by glacial
    ice until roughly 13,000 to 10,000 years ago.
 2. South of 52N, species diversity declines dramatically,  with
    a  loss  of  between  three  and  five  taxa  per  degree of
    latitude.
 3. With the exception of  those  species  able  to  persist  in
    nunataks at alpine elevations, or under arctic conditions to
    the north of the ice, or again in small, periglacial refugia
    along   the   west   coast,   most   of  British  Columbia's
    Cladoniaceae must have passed the Pleistocene south  of  the
    Cordilleran Icesheet.
 4. Floristic  and  chemical  diversity  in the Cladoniaceae are
    greater in humid regions than in arid regions, and at lower,
    forested elevations than at upper, alpine  elevations.  Many
    species can therefore be assumed to require habitats subject
    to only relatively brief periods of desiccation.
 5. Given that many Cladoniaceae probably passed the Pleistocene
    south  of  the Cordilleran Icesheet, the absence of numerous
    species from all or most of Washington, Oregon, and Califor-
    nia must reflect  climatic  changes  in  this  region  since
    deglaciation.  An  increase  in  summer moisture deficits is
    assumed to be largely responsible for this trend.
 6. Though a majority of the Cladoniaceae are  probably  now  at
    distributional  equilibrium,  a few species -- e.g., Cladina
    stellaris and C. trassii -- appear  still  to  be  extending
    their ranges southward from refugia north of the Cordilleran
    Icesheet.

References:
Geiser,  L.H.,  K.L.  Dillman, C.C. Derr & M.C. Stensvold. 1994.
   Lichens of southeastern Alaska. United States  Department  of
   Agriculture,  Forest  Service. Alaska Region, Juneau, Alaska.
   R10- TP-45.
Goward, T. & T. Ahti. 1997. Notes on the distributional  ecology
   of the Cladoniaceae (lichenized Ascomycetes) in temperate and
   boreal  western North America. Journal of the Hattori Botani-
   cal Laboratory 82: 143-155.
Hammer, S. 1995.  A  synopsis  of  the  genus  Cladonia  in  the
   northwestern United States. The Bryologist 98: 1-28.
Thomson, J.W. 1984. American Arctic Lichens 1. The macrolichens.
   Columbia University Press, New York, New York.

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