CARYOPHYLLACEAE IN NORTH AMERICA: CHANGE OVER THE LAST 110 YEARS

2008·4.1 MB·English

by  Richard K Rabeler

CARYOPHYLLACEAE NORTH AMERICA: CHANGE OVER THE LAST YEARS
IN 110
Hartman
Ronald
Richard Rabeler
K. L.
Rocky Mountain Herbarium
University ofMichigan Herbariur
Department of Botany
3600
Varsity Drive
Wyoming
Ann Michigan 48 08-2228, University of
Arbor, U.S.A.
1
Wyoming
82071-3165,
[email protected] Laramie, U.S.A.
ABSTRACT
e compare the treatment of the Caryophyllaceae in Flora of North Americ,
&
Hartman
the Caryophyllaceae in Flora North America [FNA] (Rabeler 2005) presented
of
f
Our
the opportunity to compare those results with previous efforts. treatment is the first to include all taxa
accepted North America north of Mexico since that by B.L. Robinson (1897) in Synoptical Flora of North
for
whom
FNA
The family treatment for was the combined effort of ten authors, two of also served as
Four subfamilies were recognized in which 286 species and 54 infraspecific taxa belonging to 37
editors.
genera were treated in 212 pages of illustrated text.
By contrast, the Caryophyllaceae in Synoptical Flora of North America was published solely by B.L.
Robinson. appeared as two papers (Robinson 1893, 1894) "preliminary to [the] treatment" in the
It first
Flora and then in the second fascicle of the Synoptical Flora (Robinson 1897). Three subfamilies containing
were 48 pages sans
167 species and 53 infraspecific taxa belonging to 19 genera treated in illustrations.
Bentham and
The Illecebraceae (Paronychioideae) were placed far from the Caryophyllaceae, following G.
on members
Hooker's (1862-1883) and apparently were never published. Based of this
classification,
J.D.
subfamily that were known by 1890, we estimate that 7 genera, 24 species, and 1 infraspecific taxon would
expanding
have been included in the Paronychioideae (and the tallies for the Caryophyllaceae), the treat-
ment by several pages (Table
1).
The Caryophyllaceae worldwide contain about 3000 species in 83 {Silene s.l.) or 89 (Silene s.s.) genera
&
We
which emphasizes morphological
(Rabeler Hartman 2005). followed a four-subfamily classification
features but yet to be adequately tested by molecular analyses. The four subfamilies are: Alsinoideae (30
is
200
1500 Paronychioideae genera,
genera, 1040 species), Caryophylloideae (26 [20] genera, species), (17
commonly
species), and Polycarpoideae (16 genera, 210 species). The most cited classification (Bittrich 1993)
Polycarpoideae within the Paronychioideae. Preliminary
includes three subfamilies, treating the as a tribe
and Caryophylloideae form monophyletic group within the
evidence indicates that the Alsinoideae the a
family separate from members of the other two subfamilies (Nepokroeff et al. 2002; Fior et al. 2006).
North America
The balance of this paper will compare the two treatments of Caryophyllaceae of (Rabeler
&
Hartman 2005 and Robinson 1897) by subfamily and summarize the increases in both native and alien
and
members. Our analyses take into account differences in taxonomic rank, although subspecies varieties
combined under
are the collective term, infraspecific taxa.
Table A comparison of the number of genera
1.
(g), s|
Alsinoideae
Caryophylloideae
Paronychioideae
Polycarpoideae
-se
*projected
tally
At the subfamily level, most of the differences involve the Alsinoideae (Table Changes
1). vv
subfamily (stipules absent; sepals not awned, free or only basally connate; petals usually present; fruits
capsules) encompass most of the new generic alignments and new species, both native and introduced.
Robinson followed a broad concept for Arenaria and Stellaria. The narrower concepts we employ recognized
six genera (Arenaria, Eremogone, Honckenya, Lepyrodictlis, Minuartia, and Wilhelmsia) in the former genus and
and
three (Myosoton, Pseudostellaria, Stellaria) in the latter.
By
contrast, the Caryophylloideae (stipules absent; sepals not awned, fused basally into a cup or tube;
petals present, often conspicuous; fruits capsules) treatment deviated the The major differences are
least.
Robinson's segregation of Lychnis from Silene (Morton 2005b preferred a more inclusive circumscription)
new
and
the addition of 10 native species of
Silene.
The
differences in the Paronychioideae (stipules present; sepals often awned, distinct or scarsely fused;
new
petals absent; fruits utricles) treatments involve the addition of ten taxa of Paronychia and the transfer
of Scleranihus to the Alsinoideae.
Within
the Polycarpoideae (stipules present; sepals rarely awned, distinct or scarsely fused; petals
absent or often present; fruits capsules), major changes involve the addition of the genus Polycarpaea, two
new
and
alien species of Spergula, five additional species of Drymaria.
The net change in the number of taxa between the two treatments shown in Table As noted above,
is 2.
number
the increase in the of genera (26 to 37) mostly reflects differences in generic concepts. Change in
number number
the of species (185 to 286) suggests that the of species has increased by 50 percent; this
is
somewhat some now
inflated since of Robinson's infraspecific taxa are treated as species in Cerastium).
(e.g.,
The very similar numbers of infraspecific taxa (53 54) somewhat deceiving. Besides elevation of some
vs. is
no
taxa to the species level, others are longer recognized the five varieties of Silene douglasii that Robin-
(e.g.,
On FNA
son recognized). the other hand, 22 varieties and subspecies in the treatment have been described
since 1894. Given these caveats, the total taxa comparison (species + infraspecific taxa excluding autonyms),
although increasing by about 40 percent since 1894, probably underestimates the actual change.
CHANGES
INVOLVING NATIVE TAXA
Since the beginning of North American botany, most floristic attention has been on collecting and identify-
ing our native flora. The distribution of new native species by genus shown in Table One native genus
is 3.
(Geocarpon in 1914) and 49 native species have been described since 1894. Not only are 63 percent of these
species in the Alsinoideae, but 45 percent are within Arenaria
s.l.
new
The
majority of the native species were first discovered in the western United States; not entirely
unexpected given that, prior to 1890, much of the West was poorly known botanically. In descending order,
California (12), the southwest United States (10), Texas (8), Alaska (4), and the Pacific Northwest (3) account
for 37 of the 49 species. The remaining 12 were discovered in northeastern North America
initially
(5),
southeastern United States Russia and the central United States
(2)
(3), (2).
CHANGES INVOLVING INTRODUCED TAXA
Efforts to document our alien (introduced or exotic) flora more precisely than "occurring as a weed here and
there" (Gypsophila muralis L., Gleason and Cronquist 1991) have increased in recent decades. Rabeler has
lABI
%
Robinson Rabeler&Hartman
increase
26 37 42
Species
53 54
1
Total taxa 244 340 39
Table 3. The numbers of new native species for North America by subfamily and genus.
Alsinoideae Arenaria Cerastium Eremogone Geocarpon Minua
(31): rtia(H), Pseudostellaria
(4), (5), (5), (1), (;»), Stellaria (3)
Caryophylloideae
Silene
(1 0): (1 0)
Polycarpoideae Drymaria
Spergularia
(3):
(2), (1)
been new
involved in reporting introductions of species and distribution records Caryophyllaci
for
the past 29 years (Rabeler 1980). Likewise, botanists the Rocky Mountain Herbarium
at (see Hartri
made
Nelson 2008) have a concerted effort to document invasives during more than 54 major
ii
throughout the Rocky Mountains (over 840,000 new numbered collections since 1978, both native and
in-
troduced) These initiatives long preceded the recent piqued interest in exotics. Alien taxa are very important
in the Caryophyllaceae. Of the 37 genera in the 2005 treatment, all members of 15 of those genera (41%) are
introduced to North America; this includes six of the eight genera in Caryophylloideae.
new
The distribution of introduced species by genus shown in Table Three genera (Polycarpaea,
is 4.
&
Lepyrodidis, Moenchia) did not appear in Robinson (1897) despite Moenchia erecta (L.) G. Gaertn., B. Mey.
Scherb.having been and
collected near Baltimore Philadelphia in the 1830s (Rabeler 1991). Thirty species,
them members
one-half of of the Alsinoideae, were reported since 1897; seven of these were documented
first
in North America subsequent 1960.
to
Knowing when
and where an introduction arrived in North America nearly impossible unless
first is
it was introduced intentionally. For example, Petrorhagia prolifera (L.) P.W. Ball and Heywood was brought
to North America around 1800 and appeared in John Bartram's garden in Philadelphia (Rabeler 1985). Of
may
course, the actual documentation of the plant be long introduction.
after
its
Where
documented, the first collections of "new" introductions were chiefly from coastal especially
states,
They
port areas. include the southeastern United States and Texas California the Pacific Northwest
(8), (6),
and northeastern North America Only three species appear to have been gathered in an inland
(6), (4). first
state or province (Moehringia triphylla (L.) Clairv. in Ohio, Cerastium diffusum Pers. in Missouri, and Silene
sibirica (L.) Pers. in Saskatchewan).
Some
recently discovered aliens, especially those collected in the southeastern United have
first States,
become widespread North America
in eastern Stellaria pallida (Dumort.) Crepin, Cerastium pumilum
(e.g.,
and Of
Curtis, C. brachypetalum Pers. the aliens included in Robinson (1897), some have become widespread.
now
Gypsophila paniculata L. noted as "doubtful in Manitoba" is a noxious weed there as well as in California
and Washington (Rice 1997 onwards). Holosteum umbellatum in Robinson's time known only around
L.,
now
Philadelphia, Pennsylvania, is documented from 41 states or provinces (see esp. Shinners 1965).
The
occurrence of two alien species has diminished sharply. Agrostemma githago L. and Vaccaria his-
common
panica Rauschert were formerly very wheat As
(Mill.) in agricultural practices advanced,
fields.
seed purity increased and transport of propagules of these species declined. In both cases, the range maps
FNA
in the treatments (Thieret 2005; Thieret and Rabeler 2005) are probably best described as historical
.
Journal of the Botanical Research Institute of Texas 2(2)
288
1
Table 4. The numbers of new introduced species for North America by subfamily and genus.
maximal
distributic
WHAT MIGHT THE FUTURE HOLD?
We believe that there are additional native Caryophyllaceae yet to be described in North America. Nineteen
FNA Two have been
of the 286 species in the treatment were described since 1960. additional species de-
scribed, both from California, since the 2005 treatment; Silene salmonacea T.W. Nelson, J.P. Nelson and S.A.
&
Hartman Hartman
Erwin (Nelson 2006) and Eremogone Rabeler and R.L. (Rabeler 2008).
et diftonii
al.
Likewise, the number of exotics are bound to increase. With seven species in the 2005 treatment first
dynamic more
reported since 1960, seems likely that as botanists take our alien flora seriously, the tally
it
will
rise.
In addition new discoveries and distributional records, additional study of the relationships of the
to
Many
taxa likely will change the treatment of the Caryophyllaceae. such questions are discussed in the
&
FNA treatment (Rabeler Hartman 2005) and could form the basis of interesting studies. As examples, do
(Morton 2005a) and should
and belong in other genera Silene
Stdlaria dicranoides Stellaria fontinalis really
and/or Minuartia be treated as multiple genera as discussed by Morton (2005b) and Rabeler et al. (2005)
respectively.
ACKNOWLEDGMENTS
who make comparison and Arnold Tiehm
\ treatments helped this possible
nments on an earlier draft of this paper.
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