Interspecific Hybridization and Taxon Uniformity in Arabis (Cruciferae)

Author(s): Reed C. Rollins

Source: American Journal of Botany, Vol. 70, No. 4 (Apr., 1983), pp. 625-634
Published by: Botanical Society of America
Stable URL: http://www.jstor.org/stable/2443174
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Amer. J. Bot. 70(4): 625-634. 1983.

INTERSPECIFIC HYBRIDIZATION AND TAXON UNIFORMITY

IN ARABIS (CRUCIFERAE)l

REED C. ROLLINS
Gray Herbarium, Harvard University, 22 Divinity Ave., Cambridge, Massachusetts 02138

ABSTRACT
Intermixed populations of Arabis holboellii Hornem. var. pendulocarpa (A. Nels.) Roll., A.
williamsii Roll., and their putative hybrids; and of A. holboellii var. retrofracta (Grah.) Rydb.,
A. drummondii Gray and A. divaricarpa A. Nels., the latter also considered to be a probable
hybrid taxon, were studied in the field in Montana and Wyoming. In spite of the "intermediacy"
of the putative hybrids in both cases, hybrid swarms were not found. In each instance, the
"intermediate" as well as the taxa regarded as parental were remarkably uniform. The existence
of three relatively uniform morphotypes in each of the several populations is contrary to the
usual situation involving hybridizing taxa. A possible explanation is that seed production, at
least in the "intermediate" morphotype individuals, involves apomixis, thus circumventing the
usual sexual process that would result in a hybrid swarm. Amphiploidy as a possible mechanism
for producing A. divaricarpa as a stable hybrid was ruled out in one population where the
chromosome number 2n = 14 was found in all three taxa present in the population.
Arabis divaricarpa, widely distributed in northern North America, often occurs as a uniform
morphotype where no other species of Arabis grows. If it is of hybrid origin, as the evidence
suggests, a mechanism other than sexual reproduction must be present to keep up uniformity
and at the same time perpetuate the taxon. It is suggested that facultative apomixis is involved
in the reproductive process and that it controls the genetic variability that would otherwise be
present by insuring maternal inheritance in each successive seed crop.

NATURAL INTERSPECIFIC HYBRIDIZATION is not
a rare phenomenon in the higher plants (Steb-
bins, 1950, 1969). If hybrids are local, if they
are the current product of hybridizing taxa and
reproduction is uncomplicated, the parental
species and the hybrids can easily be dealt with
taxonomically. Lesquerella provides an ex-
ample of this type (Rollins, 1957; Rollins and
Solbrig, 1973). Introgression (Anderson, 1949;
Heiser, 1949) complicates the taxonomic pic-
ture, particularly for the parent taxa. But when
polyploidy and apomixis accompany interspe-
cific hybridization and introgression, a most
complicated taxonomic pattern results (Bab-
cock and Stebbins, 1938; Rollins, 1949, 1950).
The probability that interspecific hybridiza-
tion is an important factor contributing to the
complexity of certain taxa of Arabis was early
recognized (Rollins, 1941: pp. 378-379). Since
that time, field observations on many occa-
sions (Rollins, 1946: pp. 367-369) have en-
1
Received for publication 18 March 1982; accepted 13
May 1982.
Support for research resulting in the material presented
in this paper was received from NSF Grant DEB78-08766.
I thank one of the reviewers, A. R. Kruckeberg, for con-
structive suggestions toward improvements of the paper.
hanced the view that hybridization, particu-
larly in combination with apomixis and
polyploidy (B6cher, 1951, 1969; Mulligan,
1964; Rollins, 1966) is at the base of a com-
plexity that often nearly defies taxonomic res-
olution. Particularly annoying to the taxono-
mist are those taxa where a multiplicity of
identifiable but closely related populations of
great uniformity exist, where these are readily
separable on the basis of a few characters and
where there is no easily detectable intergra-
dation between populations. If one were to for-
mally set up taxonomic entities to accom-
modate these many identifiable morphotypes,
they would become so numerous that a sensible
and utilizable classification could not be de-
veloped. Yet when such populations occur un-
der natural conditions, they cannot be ignored.
Although they present a puzzle, they must be
dealt with.
The opportunity to study combinations of
species and their intermediates in mixed pop-
ulations has presented itself in the field many
times, but only twice under favorable circum-
stances since I have been looking for an ex-
planation to the taxonomic difficulties arising
from interspecific hybridization in Arabis. The
latest opportunity came in 1979 and this has

625
626 AMERICAN JOURNAL OF BOTANY [Vol. 70

been briefly mentioned (Rollins, 1981). A fol-

low-up in 1981, together with earlier obser-
vations, provide the basis for the presentation
and discussion of Case 1.

METHODS AND OBSERVATIONS- Case I- The

taxa involved in this case are Arabis williamsii
Roll., A. holboelli Hornem. var. pendulocarpa
(A. Nels.) Roll. and their putative hybrids.
Site 1: The study of a mixed population of
Arabis species was carried out in an area about
40 ft wide and about 130 ft long on a gentle
slope between two snow fences. This site is
located near Wyoming Highway 28, 42 mi
northeast of Farson, Fremont County, Wyo-
ming, at an elevation of approximately 7,800
ft. The soils are ofgranitic origin. Sparse grasses
and scattered low sagebrush plants dominate
the area. At this site in 1979, there was a mix-
ture of the plants of Arabis holboellii var. pen-
dulocarpa, A. williamsii, and their putative hy-
brid; each was abundant. Arabis drummondii
Gray was also present at the site but it was less /! l g~~~
frequent than the other taxa. It is clear that A.
drummondii did not enter into the hybrid pic- 4cm
ture at Site 1 and no further mention of it is
made in this connection. Fig. 1. Arabis williamsii (left), R. C. & K. W. Rollins
81381; putative hybrid (center), R. C. & K. W. Rollins
In 1981, at Site 1, Arabis williamsii (Rollins, 81379; A. holboellii var. pendulocarpa (right), R. C. & K.
1981: fig. 2) occurred in great abundance and W. Rollins 81380.
the intermediate type plant was plentiful but
individuals of var. pendulocarpa were very
scarce. Only five plants could be found.
Site 2: At a location about five miles distant more tendencies towards var. pendulocarpa
from Site 1, a similar large stand of Arabis was than do the intermediates of Site 1.
found in 1981. The same taxa were present.
The three morphotypes (Fig. 1) representing Discussion of case I-A true hybrid swarm
A. williamsii, A. holboellii var. pendulocarpa, was not produced at either Site 1 or 2 as would
and their intermediates were each represented be expected if current hybridization between
by thousands of individuals but they were not the two principal taxa was actively taking place
fully intermixed. Instead, each morphotype and normal sexual reproduction was the rule.
formed pure stands in limited spots or they I emphasize that the populations at both Sites
tended to form pockets where their density was 1 and 2 were largely made up of intermixed
considerably greater than the other morpho- individuals representing the full range of mor-
types. Otherwise, there was a substantial in- phological diversity of the Arabis present, ex-
termixture of the individuals of all three mor- cluding the very different A. drummondii at
photypes. The site itself is similar to Site 1, a Site 1. Only three morphotypes were present
gentle slope on the lee side of a snow fence. at each site and each of these was relatively
At Site 2, the morphotype referable to Arabis uniform. One hypothesis to explain the un-
williamsii is not exactly the same as that of usual and unexpected situation would be that
Site 1. The plants of the population are uniform the three discrete and different morphotypes
but the pedicels and siliques diverge slightly represent independent taxa. If this were the
from the rachis of the infructescence (Fig. 1), case the intermediate type would need to be
rather than being strictly erect as in plants of established in the literature as a non-hybrid
Site 1. In addition, the Site 2 plants differ in taxon in its own right. However, it is difficult
having pubescence that carries up the stem to visualize how such a taxon could have arisen
more definitely than in those of Site 1. Also, without involving the hybridization of the two
the intermediates at Site 2 are not exactly like most divergent morphotypes with which it is
the intermediates at Site 1. They show a few associated. The intermediates do not have dis-
April, 1983] ROLLINS-INTERSPECIFIC
tinctive features that cannot be explained by
merely assuming a recombination of the fea-
tures of the putative parental taxa. So far, the
intermediates have been found only in asso-
ciation with the taxa from which they are pre-
sumed to have arisen by hybridization.
The hypothesis that the intermediates are of
hybrid origin requires the assumption that a
breeding system other than normal sexuality
is operative. If normal sexuality were predom-
inant, a hybrid swarm in which more or less
of a continuum of variation between the ex-
treme morphotypes would be present. How-
ever, if hybridization were followed by apo-
mixis, particularly if the hybridization was
ancient, then the situation found in the mixed
populations of both sites is explainable. This
hypothesis is testable but the resources to do
this are not presently available to me. But apo-
mixis is known in a number ofNorth American
Arabis species (B6cher, 1969) and its presence
as part of the reproductive apparatus in this
case would not only explain the uniformity of
the intermediate morphotypes, it could be in-
voked to explain the presence of slightly dif-
ferent morphotypes of Arabis williamsii be-
tween Sites 1 and 2 (compare left plant of Fig.
1 and Rollins, 1981: fig. 2). At Site 2, it is
probable that some introgression, secondarily
involving var. pendulocarpa, took place and
the resulting introgression became fixed
through apomixis. This would account for the
presence of somewhat different intermediates
at Site 2 and Site 1. If the slight differences of
A. williamsii at Sites 1 and 2 are also due to
introgression followed by apomixis, this would
also account for the uniformity of these two
slightly different morphotypes in their respec-
tive localities.
Alternative hypotheses to explain the situ-
ation in the populations of Sites 1 and 2 are
less plausible than that involving hybridity
plus apomixis. For example, it is difficult to
see how selective pressure could be operative
in such a way as to bring about and preserve
three distinct morphotypes, represented by
hundreds to thousands of individuals, with one
exception, in each of the sites studied. All in-
dividuals are subject to the same pressures and
the habitat is sufficiently uniform to preclude
local sorting on the basis of edaphic factors.
As another alternative, it is scarcely conceiv-
able that all of the intermediates, again
amounting to thousands of individuals, could
be F1 hybrids. If the crossability of Arabis hol-
boelliivar.pendulocarpaandA. williamsiiwere
as facile as would be necessary for that, the
probability of backcrossing and the ultimate
HYBRIDIZATION IN ARABIS 627
production of a typical hybrid swarm would
seem to be inevitable.
The possibility that inbreeding could com-
bine with hybridization to produce the situa-
tion found should also be considered. It is quite
probable that all three morphotypes are largely
autogamous and could be inbreeding although
I have no evidence for this. My experience with
natural inbreeding populations of Leavenwor-
thia, another genus of the Cruciferae, is that
they tend to form on the basis of a single mor-
photype (Lloyd, 1965; Rollins, 1963). The
population profiles at both Sites 1 and 2 do
not accord with expectations if inbreeding were
the controlling genetic mechanism.
From this study and that of B6cher cited
above, it is clear that both interspecific hy-
bridization and apomictic reproduction are
factors that must be taken into account in de-
termining the taxonomy of Arabis where A.
holboellii var. pendulocarpa is involved. Ap-
plying what we understand from the popula-
tions of Sites 1 and 2, it now is more certain
than tentatively suggested previously (Rollins,
1981) that A. microphyllavar. saximontana is
of hybrid origin. Since A. williamsii is not yet
known from the areas in Idaho and Wyoming
where var. saximontana grows, one cannot be
sure that is the species involved with var. pen-
dulocarpa to produce var. saximontana. But
if the latter is apomictic as suspected, it could
have moved to its present location indepen-
dent of its putative parental taxa. However, it
is more likely to have arisen independently at
the several different places from which it is
known. Regardless of its mode of origin, var.
saximontana is taxonomically distinctive and
requires nomenclatural recognition, and it
should be associated with A. williamsii, not A.
microphylla.2
Case 2-The three taxa involved are Arabis
drummondii Gray (2n = 14), A. holboelli Hor-
nem. var. retrofracta (Grah.) Rydb. (2n = 14)
and A. divaricarpa A. Nels. (2n = 14). These
were growing together at three different sites,
two in Park County, Wyoming, and one near
Cooke, Park County, Montana. Both basal ro-
settes by themselves and mature plants of all
three taxa were growing intermixed at all three
sites. The first-year basal rosettes of these three
taxa are easily distinguished from each other
(Fig. 2). The leaves of A. drummondii are strig-
illose with malpighiaceous trichomes; those of
A. holboellii var. retrofracta are densely matted
2Arabis williamsii Roll. var. saximontana (Roll.) Roll.,
comb. nov., based on A. microphylla Nutt. var. saximon-
tana Rollins, Rhodora 43: 429. 1941.
628 AMERICAN
Fig. 2. Leaf-surfaces of Arabis showing trichomes. X20. A. drummondii (left), A. divaricarpa (middle), A. holboell/i
var. retrofracta (right).
with fine dendritically branched trichomes;
and those of A. divaricarpa are covered with
3- to 5- or 6-branched trichomes. In A. drum-
mondii, the siliques are strictly erect; in var.
retrofracta, the siliques are strictly reflexed; and
in A. divaricarpa the siliques are divaricately
ascending to very widely descending. It will be
noted that in these features, A. divaricarpa is
more or less intermediate between A. drum-
mondii and A. holboellii var. retrofracta (Fig.
3). This is true for other characteristics as well.
For example, the seeds of A. drummondii are
about half the silique width in size and they
JOURNAL OF BOTANY [Vol. 70
are arranged in two definite rows extending the
length of the siliques. The seeds of A. holboelli
var. retrofracta are larger and fill the full width
of the siliques. These are arranged in a single
row. The seeds of A. divaricarpa are inter-
mediate as to size and they alternate on one
side, then the other, forming an imperfect row
that is neither wholly single nor wholly double.
The malpighiaceous trichomes of A. drum-
mondii are relatively large with the two branch-
es closely appressed to the leaf surface. In A.
holboellii var. retrofracta, the dendritic tri-
chomes are minute and although densely mat-
April, 1983] ROLLINS-INTERSPECIFIC HYBRIDIZATION IN ARABIS 629

30

' 20
0

6
C 0

0 1 2 3 4 5 6 7 8 9
Population 51250 N:121

30

20
4.5 cm

Fig. 3. Arabis drummondii (left), Rollins 51246; A. di-

varicarpa (middle), Rollins 51244; A. holboellii var. ret-
0
rofracta (right), Rollins 51245.

ted they have a central stalk elevating the

branches above the leaf-surface. The trichomes
of A. divaricarpa are somewhat in between,
neither as large as those of A. drummondii nor
as small as those of var. retrofracta, and the 3
to 5 or 6 branches are somewhat elevated on
a central stalk.
Randomly gathered rosettes from two pop-
ulations were scored for trichome character-
istics in the following way: 0 = trichomes
wholly malpighiaceous and closely appressed;
1 = trichomes mostly malpighiaceous, with a
few 3-parted; 2 = trichomes mostly 3-parted,
with a few malpighiaceous; 3 = trichomes
3-parted; 4 = trichomes 3- and 4-parted; 5 =
trichomes with 4 and 5 branches; 6 = tri-
chomes with 5 and 6 branches; 7 = trichomes
with 6 and 7 branches; 8 = trichomes with
many branches but coarser than in 9; 9 = tri-
chomes fine, matted, and with many branches.
Figure 4 gives the results from population
51250, growing near Clarks Fork River, 9 mi
southeast of Cooke, Montana, in extreme
northwestern Park County, Wyoming; and
population 51263, growing at 7,500 ft eleva-
tion in a rocky opening of a spruce-fir forest,
near U.S. Hwy. 12, one-half mi south of the
0 1 2 3 4 5 6 7 8 9
Population 51263 N=121
Fig. 4. Graphs from index scores of trichome-branch-
ing on first year rosettes of two intermixed populations of
Arabisdrummondii,A. divaricarpaand A. holboelliivar.
retrofracta. Each graph is based on a sample of 121 rosettes.
See text for explanation of index numbers.
Montana state line in extreme northwestern
Park County, Wyoming. Zero and one are Ar-
abis drummondii, 9 and 8 are A. holboellii var.
retrofracta, and 3-7 are A. divaricarpa (Fig. 4).
It is clear from the analysis of rosettes that
there is great uniformity in the trichomes of
Arabis drummondii and in A. holboellii var.
retrofracta but there is a considerable range of
variation in A. divaricarpa. The same holds
true for silique position in these populations.
In A. drummondii, the siliques were uniformly
erect; in var. retrofracta they were uniformly
reflexed; but in A. divaricarpa, the silique po-
sition varied from divaricately ascending to
loosely pendulous. These characters and others
show A. divaricarpa to be intermediate be-
630 AMERICAN JOURNAL
tween A. drummondii and A. holboellii var. stone conglomerate, near golf course, Bic, 29
retrofracta. Aug. 1955, Rollins & Wood 55242. A micro-
The evidence, taken together, suggests that
Arabis divaricarpa is of hybrid origin. Such a
hypothesis fits with the known facts. It also
seems probable that it is largely apomictic. As
it is presently understood, A. divaricarpa may
have arisen many times in many different
places. Is polypoidy and possibly aneuploidy
also involved? Bocher (1954) grew what ap-
peared to be hybrids of A. divaricarpa X hol-
boellii with 2n = 22. Mulligan (1964) exam-
ined material of A. divaricarpa from 13
localities in Canada and reported chromosome
numbers of 2n = 13 + 2B, 2n = 14, 2n = 20,
2n = 20 + 2B, and 2n = 21. Among five dif-
ferent populations, I found numbers of 2n =
14 and 2n = 22 (Rollins, 1966). Assuming that
x = 7 is the fundamental number for the taxon,
it is clear that polyploidy is one factor involved
in producing the polymorphy present in A. di-
varicarpa. Aneuploidy and the presence of su-
pernumerary B chromosomes may also con-
tribute to the overall diversity present in the
species.
Additionalfield observationson Arabis di-
varicarpa-Site 1: Quebec, Canada, Rimouski
Co.: near the mouth of Riviere Hatee, 2 mi
north of Bic, 28 Aug. 1955, Rollins & Wood
55228. The numerous plants of this population
were growing among slaty rocks. Basal rosettes
and mature individuals were present. The tri-
chomes on 129 rosettes were examined. These
were typically 3-parted and the branches were
elevated on a stalk above the leaf-surface. Both
4-branched and a few 5-branched trichomes
were sometimes found mixed with those with
3 branches. Leaf-shape was about the same in
all rosettes except those growing in the shade
when the leaves were considerably larger and
the trichomes spaced away from each other.
There were no other species of Arabis at the
site and there was no evidence of hybridization
or introgression.
Site 2: Quebec, Canada, Rimouski Co.:
coastal rocks, southeast and eastern side of Ilet
a d'Amours, ca. 1 mi west of Bic, 29 Aug. 1955,
Rollins & Wood 55235. Two of the 16 rosettes
had a few bipartite trichomes suggesting some
possible introgression from Arabis drummon-
dii but the other 14 had 3-parted or occasion-
ally 4-parted trichomes. Arabis holboellii Hor-
nem. var. collinsii (Fern.) Roll. grew in the
vicinity but no evidence of its involvement in
hybridization could be found. Fifty-one ro-
settes of var. collinsii were examined in this
connection.
Site 3: Quebec, Canada, Rimouski Co.: lime-
OF BOTANY [Vol. 70
scopic examination of 15 rosettes showed no
evidence of introgression from Arabis hol-
boelli var. collinsii although the latter was even
more abundant than A. divaricarpa at the site.
The basal rosettes of the two taxa were easily
distinguished from each other and a careful
examination of 42 rosettes of var. collinsii did
not reveal any evidence of contamination from
A. divaricarpa. There was a slight ecological
preference shown by the two taxa: Arabis di-
varicarpa tended to be in slightly wetter places
than var. collinsii.
Site 4: Quebec, Canada, Rimouski Col: grav-
el and small rocks, southern base of west end
of ridge, south of Cap Enrage, ca. 4 mi west of
Bic, 29 Aug. 1955, Rollins & Wood 55247.
There is a little more variation in the trichomes
of the 20 rosettes examined from this site than
from the other three Rimouski County sites.
But there are no definite trends and no evidence
ofintrogression from other species even though
Arabis holboellii var. collinsii was present at
the site.
Site 5: Wyoming, Fremont Co.: 13 mi north
of Dubois toward Ramshorn Peak, 8,400 ft
elevation, 27 Jul. 195 1, Rollins & Porter
51242. Silique position varied from spreading
at right angles to the rachis ofthe infructescence
to loosely descending. Trichome type on the
basal leaves was roughly correlated with silique
position. Trichomes on plants with widely
spreading siliques were 3- to 4-branched.
Those on plants with descending siliques were
somewhat dendritic with five or more branch-
es.
Site 6: Idaho, Lemhi Co.: near Eight Mile
Creek, base of Gunsight Peak, 7,500 ft ele-
vation, 22 Jul. 1957, Rollins 57247. Tri-
chomes on 18 rosettes examined were mostly
3-branched with some 4-branched and a few
5-branched. The siliques were loosely descend-
ing. Both mature plants and rosettes were rea-
sonably uniform and showed no evidence of
recent hybridization with Arabis holboellii var.
retrofractaor A. microphyllaNutt. var. ma-
counii (S. Wats.) Roll., both of which were in
the same locality.
Site 7: Idaho, Bonneville Co.: meadow near
Rainey Creek, 8.3 mi northeast of Swan Val-
ley, 20 Jun. 1979, R. C. & K. W. Rollins 79296,
79297, 79298, 79299. Samples were selected
on the basis of silique position; 79296, strongly
ascending but not strictly erect; 79297, divar-
icately ascending to more widely spreading;
79298, widely spreading to loosely descending;
79299, strongly descending. This large popu-
lation looked like a hybrid swarm representing
April, 1983] ROLLINS-INTERSPECIFIC
intermediates in nearly all combinations be-
tween Arabis drummondii on the one hand and
A. holboellii var. retrofracta on the other. How-
ever, neither A. drummondii nor var. retro-
fracta were present at the site. The trichome
branching roughly correlated with silique po-
sition and ranged from the most A. drum-
mondii-like in 79296 to the most var. retro-
fracta-like in 79299. Since this population was
in a meadow which could be flooded from time
to time by Rainey Creek, it is possible that the
seeds were transported from a site where A.
drummondii and A. holboellii var. retrofracta
hybridized to produce seeds ultimately giving
rise to the population we sampled.
Site 8: Idaho, Bear Lake Co.: steep bank in
a spruce-aspen area, Paris Canyon road, 6.7
mi west of Paris, 16 Jun. 1981, R. C. & K W.
Rollins 81324. This population of Arabis di-
varicarpa was notable because the plants were
very large, up to 5 ft tall. Usually plants of A.
divaricarpa are 1 to 2 ft, rarely up to 3 ft in
height. Could hybrid vigor be involved here?
There were no other species of Arabis present.
Site 9: Colorado, Park Co.: disturbed area
along roadside, one mile south of Hoosier Pass,
10,800 ft elevation, 7 Aug. 1951, Rollins
51290. Both Arabis divaricarpa and A. drum-
mondii were present. The siliques of A. divar-
icarpa were uniformly straight and divaricately
ascending. Trichomes were few on the basal
leaves but those present were either malpigh-
iaceous and appressed along the petiole mar-
gins or bifurcate and stalked. None were ap-
pressed as in A. drummondii. The best
hypothesis is that A. divaricarpa at this site was
an apomictic line originating from a back-
crossing situation where A. drummondii was
predominant.
Site 10: Utah, Daggett Co.: aspen-pine as-
sociation near Sheep Creek, 12 mi south of
Manila, ca. 7,000 ft elevation, 27 Jun. 1951,
Rollins & Porter 5130. Different plants in this
population referable to Arabis divaricarpa had
different silique positions. These ranged from
divaricately ascending through widely spread-
ing to loosely pendant. The complexity of the
trichomes as indicated by the number of
branches followed the same sequence; the sim-
plest trichomes were correlated with the more
nearly erect -siliques and the most highly
branched trichomes with descending siliques.
There were no other species of Arabis in the
immediate vicinity.
Site 11: Utah, Uintah Co.: dry knoll, ca. 25
mi north of Vernal, 8,000 ft elevation, 26
Jun. 1951, Rollins & Porter 5123. The siliques
of the plants were uniformly pendant from
arched pedicels on all plants determined as
HYBRIDIZATION IN ARABIS 631
Arabis divaricarpa. On these plants the tri-
chomes had 4 to 5 branches and were about
the same on all samples studied. Both Arabis
drummondiiand A. holboellii var. retrofracta
were present at the site. The total Arabis pop-
ulation was characterized by three morpho-
types only, each distinct from the others and
clearly representing a different taxon.
Site 12: Utah, Box Elder Co.: sagebrush area,
10.9 mi north of Grouse Creek on road to Lynn,
9 Jun. 1981, R. C. & K. W. Rollins 81257. A
few plants were found with divaricately as-
cending siliques and 3- or 4-branched tri-
chomes, but most of the population consisted
of plants with widely spreading to slightly pen-
dant siliques and trichomes with 4-6 branches.
Arabis holboellivar. retrofractaand A. drum-
mondii were in the same vicinity.
Discussion of case 2-Arabis divaricarpais
highly polymorphic compared to its close rel-
atives Arabisholboelliis.l. and A. drummondii.
In some places, the populations of it are rel-
atively uniform; in others what appear to be
several recombinant types of hybrids occur
consisting of some individuals clustering near
A. drummondii,others near A. holboellii, but
with more nearly median morphotypes usually
predominating. In those instances where a sin-
gle morphotype of A. divaricarpa is present it
may be about midway between the putative
parental taxa as in the populations in the vi-
cinity of Bic, Quebec, as described above, or
nearer to A. drummondii, as in the population
from near Hoosier Pass, Colorado, or closer to
A. holboelli as in the population described
from Uintah County, Utah. Given hybridiza-
tion accompanied by apomixis, uniformity can
be achieved at any position on the spectrum
of differences between the hybridizing taxa.
This is apparently what has happened in A.
divaricarpa. The taxon as we know it consists
of ancient hybrids, in many cases narrowed by
selective pressures and apomictic reproduction
to a uniform morphotype; recent hybrids
where a range of morphotypes persist in a given
population; and many intermediate situations.
A true hybrid swarm, where both parental taxa
and the hybrids are totally intermixed, has not
been seen. This unusual situation is attributed
to the probability that normal sexual repro-
duction does not predominate in most popu-
lations either in the parental taxa or the hy-
brids. It is likely that both sexuality and
facultative apomixis are a part of the repro-
ductive pattern of A. drummondii, and A. hol-
boelliivar. retrofractaas well as A. divaricarpa.
The possibility that amphiploidy is involved
was investigated in plants grown from the three
632 AMERICAN JOURNAL OF BOTANY [Vol. 70

sites in Park counties, Montana and Wyoming. phyletic in origin. There are many species with
The need for this was suggested particularly by disjunct distributions that have a disrupted di-
population 51250 where the putative hybrids versity similar to the situation in A. divaricar-
were almost exactly intermediate between Ar- pa. When this is the case, they could hardly be
abis drummondiiand A. holboelli var. retro- described as having a close-knit morphological
fracta. Also, the hybrids were relatively uni- integrity. But they are presumed to be mono-
form. The chromosome number of all three phyletic and when there is no evidence to the
taxa from all three sites was 2n = 14, thus am- contrary, they are regarded as species.
phiploidy was eliminated as a possible mech-
anism of producing stable intermediate hy- GENERAL DISCUSSION- The name Arabis di-
brids in at least these three populations. varicarpa-Use of the name Arabis confinis S.
A taxonomic and philosophical question is Wats. instead of A. divaricarpa A. Nels. has
raised by the nature of Arabis divaricarpa as been advocated by Welsh and Reveal (1977).
presently understood. Can a species exist that Earlier Fernald especially (1903) and later
is more or less continuously originating? Ap- Hopkins (1937) dealt with the problem of ac-
plying the axiom that it matters not how a cepting the older A. confinis which was pro-
species originates, by mutation or by hybrid- posed by Watson (1887) to "include all of the
ization for example, what matters to the tax- 'A. drummondii' of the Atlantic region." Con-
onomist is what happens after the events of trary to the statement of Welsh and Reveal
origination have occurred. Are new characters that A. confinis was based on Turritis brachy-
or combinations of characters produced that carpa T. & G., it was in fact based on the
persist? Are populations that persist in time treatment of A. /aevigata by Hooker (1 840). A
achieved? Is there an integrity in space as well specimen collected by Dr. Todd about Lake
as time that holds the taxon together? What Huron is the only one cited by Hooker and it
kind of variation is tolerable if a taxon is to automatically becomes the type of A. confinis.
be called a species? Does the variation have to I have not seen the Todd specimen which
be of a continuous nature or can it be disjunct should be in the herbarium at Kew. However,
and erratic? If interspecific hybridization is the Hooker's short description of the collection fits
primary mode of origin for A. divaricarpa, what A. drummondii better than it does A. divari-
range of participating taxa is permissible if a carpa. Therefore, it seems best to continue the
single hybrid taxon is considered to have been use of A. divaricarpa at least until the Todd
produced? For example, in Quebec it is A. hol- specimen can be examined to see whether it
boellii var. collinsii that apparently has hy- is, as I suspect, A. drummondii or whether it
bridized with A. drummondii to give rise to A. is the same as A. divaricarpa. If it is the latter,
divaricarpa; in the Rocky Mountain area, it is then A. confinis must be adopted.
A. holboelliivar. retrofractaX A. drummondii. The bearing of these findings on Arabisfruc-
Some authors (Boivin, 1951; Hopkins, 1937) ticosa: A special interest in Arabis fructicosa
treat either var. collinsii or var. retrofracta or arises from the facts that it has not been re-
both as distinct species. If we demand that a collected since the original material was gath-
species be strictly monophyletic, then A. di- ered in 1899, that the type material was from
varicarpa does not qualify as a species. Arabis Yellowstone Park, a government preserve, and
divaricarpa does not qualify as a hybrid taxon that it has been listed as an endangered species
in the strict sense either. Although we may by government agencies. I have searched in the
suggest that distinctive populations, remote type locality for Arabis that would be referable
from either or both of the putative parental to A. fructicosa on two separate occasions and
species, are of hybrid origin, this cannot be several others have combed the type area many
proved. It should be stressed that in many times. None of us have found an Arabis that
places A. divaricarpa exists by itself without matches the type of A. fructicosa. The nearest
any contact with other taxa of Arabis. approach among specimens I have studied is
Many of the criteria we would apply, to de- a collection from Park County, Montana, with
termine whether or not Arabis divaricarpa is a the following data: slopes of Woodie Mountain
species in its own right, are fully met. New and facing Republic Creek, a mile or two south of
distinctive characters and character combi- Cooke City, Aug. 2, 1949, Jean G. Witt 1869
nations are present and they do persist over (GH). Specimens of this collection I would re-
time. Populations are established that persist fer to A. divaricarpa. A second collection from
for many years. Where A. divaricarpa fails to Fremont County, Wyoming, Dickinson Park,
meet certain species criteria is in its lack of meadows along Dickinson Creek, 9,300 ft el-
morphological integrity coupled with the fact evation, June 24, 1965, Richard W. Scott 455
that the diversity present appears to be poly- (GH), has some features similar to those of A.
April, 1983] ROLLINS- INTERSPECIFIC
fructicosa but this, too, I would determine as
A. divaricarpa. A third collection, distributed
as A. holboellii var. pendulocarpa, C. L. & M.
W. Porter 10438 (GH) from Sublette County,
Wyoming, may be able to qualify as A. fruc-
ticosa when the nature of that putative taxon
is better known.
At this point, I suggest as a possibility that
the type material of Arabisfructicosa was pro-
duced as a segregate from the hybrids of A.
drummondii and A. holboelli var. retrofracta
in which some characters are combined in a
unique way scarcely matched by other known
descendants of this cross. If that were the case,
A. fructicosa as such, with its particular set of
characteristics, may not have persisted as a
population. The original habitat was given as
"on dry roadsides" which probably meant a
disturbed site. We know that disturbed sites
are the right place to look for hybrids. The
reason A. fructicosa has not been rediscovered
in over 80 years may be that the right com-
bination of characters in a hybrid population
has not been available when the searchers have
been in the area. It might be suggested that the
right combination ought to have turned up
somewhere in the vast geographic area where
A. drummondii and A. holboellii var. retro-
fracta are sympatric. Indeed, this is an argu-
ment against the hypothesis of hybrid origin
but it should be remembered that sampling of
the thousands of square miles involved is very
thin.
I do not wish to go on record at this time as
being totally committed to the notion that Ar-
abis fructicosa does not exist as a species. All
of the questions concerning it have not been
answered. But it is possible that the reason it
has not been rediscovered is that it was never
there at the beginning as a species independent
of all others. Thus strenuous efforts to redis-
cover A. fructicosa may not be easily justified
at the present time. Over time with normal
floristic inquiry the situation is likely to be
clarified.
Conclusions-Relatively uniform popula-
tions that appear to have arisen from inter-
specific hybridization between Arabis hol-
boelli s.l. and A. drummondii and between A.
holboelli var. pendulocarpa and A. williamsii
call for interpretations inconsistent with the
usual picture of hybridization in higher plants.
Three aspects are unusual: the lack of a hybrid
swarm; the uniformity of the "intermediate"
morphotype in given populations; and the fre-
quent occurrence of the A. divaricarpa "inter-
mediate" by itself in populations of thousands
of individuals without either of the putative
parental taxa being present. The presence of
HYBRIDIZATION IN ARABIS 633
three distinct morphotypes in intermixed pop-
ulations where hybrids occur but a hybrid
swarm is not produced is explainable if apo-
mictic seed production dominates the repro-
ductive pattern in at least the "intermediate"
morphotype. If apomictic seed production pre-
dominates in the putative parental taxa as well,
the maintenance of the relatively uniform mor-
photypes in intermixed populations in a sit-
uation where some but not much hybridization
also occurs would be even more effectively
achieved. If an "intermediate" morphotype
could maintain itself in an intermixed popu-
lation, where it is theoretically subject to being
"hybridized" out of existence, there is no ob-
stacle to its maintaining itself outside of the
intermixed population. This possibility would
account for the occurrence of A. divaricarpa by
itself in many locations remote from either or
both of its putative parental taxa. Similarly, it
would explain why A. williamsii var. saxi-
montana persists in a number of localities out
of contact with possible progenitors for pro-
ducing it by hybridization.
The taxonomic interpretation of Arabis di-
varicarpa, in view of its probable hybrid origin,
can go in either of two directions. It can be
considered to be a natural hybrid taxon rec-
ognizing that as now treated it includes stable
populations of ancient hybrid origin, less stable
more recently produced hybrids, and many hy-
brid populations produced under intermediate
conditions. Furthermore, the array of mor-
photypes collectively included in A. divaricar-
pa range from near A. holboellii s.l. to near A.
drummondii but with most populations show-
ing a high degree of uniformity. Perhaps the
most frequent morphotype encountered is
roughly intermediate between the presumed
parental taxa. An alternative interpretation is
the one followed for many years in which A.
divaricarpa has been treated as a species. If this
interpretation is followed, it must be realized
that what is now called A. divaricarpa is a
polytypic species and there are those who do
not admit that such a species can exist. If it
were possible to narrow down the inclusiveness
now recognized in such a way as to eliminate
the more obvious hybrids, A. divaricarpa
might become a more integrated taxon mor-
phologically but the problem of where to place
the hybrid types thus excluded would remain.
In the taxonomic system, there is no large ad-
vantage to recognizing A. divaricarpa as a hy-
brid taxon and I prefer to follow tradition and
treat it as a species but with the full recognition
that it is probably of hybrid origin.
The case of Arabis holboelli var. pendulo-
carpa X A. williamsii is somewhat different
634 AMERICAN JOURNAL
from that of A. divaricarpa. Instead of being a
widespread species spanning the continent, as
in A. divaricarpa, this hybrid situation appears
to be much more local and the consequences
of interspecific hybridization much less well
worked out. For the present, the hybrid com-
bination is sufficient to indicate what has hap-
pened in the two known populations. Other-
wise, A. williamsii var. saximontana
is
recognized as possibly being of hybrid origin
but any further treatment of it as such awaits
further evidence.
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70