The 2001 Bryophyte Expedition on the Santa
Margarita Ecological Reserve,
in the Peninsular Ranges of Southern California
Steven L. Jessup[1],
Ken Kellman[2],
John Spence[3],
James R. Shevock[4],
Lloyd R. Stark[5],
William Doyle[6]
Abstract. Bryophytes collected on the Santa Margarita Ecological Reserve
in the coastal peninsular Santa Margarita Mountains of Southern California
are documented. Collections were made during the sixth
annual Spring Outing, Botanical Excursion, Foray, Retreat, and Escape to
the Environment (SO BE FREE), and include 65 moss taxa in 32 genera and 12
families, 12 liverwort taxa in 7 genera and 6 families,
and 2 hornwort taxa. Collections included plants representing
3 new species in Bryum. New localities
were discovered for the rare and little known bryophytes, Bryum
pyriferum, Schizymenium shevockii, Tortula californica,and Sphaeropheros drewei.
Introduction
The Santa
Margarita Ecological Reserve is centered around Temecula Canyon, at about
33°27'N, 117°11'W, on the San Diego County - Riverside County line (fig.
1). The reserve, one of four San Diego State University
field stations, occupies 4344 acres, ranging in elevation from 150 to 700
m, situated approximately 30 km from the Pacific Coast in the Santa Margarita
Mountains at the southern extreme of the Santa Ana Range.
The vegetation is a mosaic of coastal sage scrub, oak woodlands, and
chapparal. The steep and rolling terrain is punctuated by cliffs and riparian
corridors. Annual precipation averages about 400 mm
with most of the precipitation occuring during the relatively cool winters. Mean annual temperature is approximately 16.4°C.
The Sixth
Annual Meeting of Spring Outing, Botanical Excursion, Foray, Retreat, and
Escape to the Environment (SO BE FREE), convened at the Santa Margarita Ecological
Reserve Field Station 24-27 March 2001. During the
meeting the attentions of the more than thirty botanists attending were focused
on compiling a list of taxa in the bryophyte flora of the reserve. This report summarizes the findings of that effort with
a list of taxa comprising approximately 12% of the known bryophyte flora of
California, including 3 taxa new to science.
On the first day of the foray we explored along the Santa Margarita River
in Temecula Canyon. This riparian area is subject to periodic flooding and
scouring. Proximity of the city of Temecula with its polluted runoff, and
other upstream uses contributing to eutrophication, such as agriculture, adversely
affects the aquatic bryophyte component of the river corridor. Pools and river margins exhibited a lush development of
fresh-water algae, probably due to nitrogen and phosphorus inputs, apparently
at the expense of bryophytes since very few taxa were found in the river.
Sandy soils comprise the flood zone while the canyon walls are a mixture of
granitic and metasedimentary rocks within a dense woodland and chaparral. Excursions on the second day focused on the upland portions
of the reserve, primarily in the California live oak woodlands and associated
openings in chamise chaparral above the river canyon.
The bryophyte flora of the southwestern U.S. and northwestern Mexico has
been previously documented in a series of papers (Bowers, Delgadillo &
Sharp, 1976; Harthill, Long & Mishler, 1979; Stark & Whittemore,
1992) and in a recent flora (Sharp, Crum & Eckel, 1994). That considerable bryophyte diversity remains to be discovered
in this region is clearly demonstrated by the series of novelties and rarities
discussed below. Collection numbers in the following
list are indicated with a three-letter prefix: DYL=Doyle, JSS=Jessup, KLL=Kellman,
SHV=Shevock, SPC=Spence. Shevock and Kellman collections
are housed at CAS. Doyle and Spence collections are
in their personal collections. Jessup collections are
housed at SOC.
Nomenclature
generally follows that of Anderson, Crum and Buck (1990), and Zander (1993)
for the Pottiaceae. Family designations follow Buck
& Goffinet (2000).
Hornworts
Anthocerotaceae
Anthoceros
fusiformis Aust. [DYL 9898, 9909]
Phaeoceros
pearsonii (Howe) Prosk. [DYL 9911; 13-62]
Liverworts
Aytoniaceae
Asterella
californica (Hampe) Underw. [DYL 9902; JSS 13-08]
palmeri (Aust.) Underw. [DYL 9901; JSS 13-03]
Cryptomitrium
tenerum (Hook.) Aust. [DYL 9901; JSS 13-33]
Cephaloziellaceae
Cephaloziella
byssacea (Roth) Warnst. [DYL 9903, 9907; JSS 13-15, 13-30, 13-52]
Fossombroniaceae
Fossombronia
longiseta Aust. [DYL 9899; JSS 13-04; SHV 20516]
Ricciaceae
Riccia
campbelliana Howe [DYL 9895; JSS 13-11]
nigrella DC [DYL 9896; JSS 13-05]
sorocarpa Bisch. [DYL 9894; JSS 13-17]
trichocarpa Howe [DYL 9897; JSS 13-01]
Sphaerocarpaceae
Sphaerocarpos
drewei Wigglesw. [JSS 13-68]
texanus Aust. [DYL 9898, 9909]
Targioniaceae
Targionia
hypophylla L. [DYL 9900; JSS 13-02, 13-16, 13-41]
Mosses
Bartramiaceae
Anacolia
menziesii (Turn.)Par.
[SHV 20485]
Bartramia
stricta Brid. [JSS 13-54; SHV 20534]
Brachytheciaceae
Eurynchium
hians (Hedw.)
Sande-Lac. [SPC 5430]
Homalothcium
arenarium (Lesq.) Lawt. [SHV 20486]
Scleropodium
cespitans (C. Müll.) Koch [KLL 1380]
colpophyllum (Sull.) Grout [JSS 13-60; SHV 20450, 20509]
touretii (Brid.) L.Koch [JSS 13-32]
Bryaceae
Anomobryum
argenteum (Hedw.) Spence [SHV 20489; SPC 5434]
lanatum (P. Beauv.) Spence & Ramsay [KLL xxx]
Bryum
bicolor Dickson [SHV 20491, 20513; SPC 5432; KLL 1366]
caespiticium Hedw. [KLL 1379; SPC 5439]
dichotomum Hedw. [KLL 1366]
gemmilucens Wilcz. & Dem. [JSS 13-37, 13-42; KLL 1388; SPC 5433]
gemmiparum De Not. [KLL 1382, 1383]
pseudotriquetrum (Hedw.)
Gaertn., Meyer & Scherb. [SHV 20515]
pyriferum Crund. & Whiteh. [SHV 20496; SPC 5435]
species nov. B [KLL xxxx]
species nov. C [KLL 1374a]
species nov. D [KLL xxxx; SPC 5429]
Rosulabryum
capillare (Hedw.) Spence [SPC 5437]
Ditrichaceae
Ceratodon
purpureus (Hedw.) Brid. [KLL 1372]
Pleuridium
acuminatum Lindb. [JSS 13-06; KLL 1367; SHV 20488]
Fissidentaceae
Fissidens
sublimbatus Grout [JSS 13-18, 13-27;
KLL 1364; SHV 20483, 20499]
Funariaceae
Funaria
hygrometrica Hedw. [JSS 13-09; SHV 20493]
muhlenbergii Turn. [KLL 1384, 1387]
Grimmiaceae
Grimmia
laevigata (Brid.) Brid. [JSS 13-38, 13-58; KLL 1397; SHV 20507]
lisae DeNot.
[KLL 1398]
montana B.S.G. [KLL 1377]
pulvinata (Hedw.) Sm. ex Sm & Sowerb. [JSS 13-36; SHV 20487]
reflexidens C. Müll. [JSS 13-43]
trichophylla Grev. [SHV 20484]
ungeri Jur. [KLL 1373]
Schistidium
cinclinodonteum (C. Müll. in Röll.) Bremer [JSS 13-28]
Leskeaceae
Claopodium
whippleanum (Sull.) Ren. & Chard. [JSS 13-59]
Mniaceae
Schizymenium
shevockii J. Shaw [SHV 20506]
Orthotrichaceae
Orthotrichum
tenellum Bruch ex Brid. [JSS 13-61, 13-64, 13-65; KLL 1389]
Pottiaceae
Aloina
aloides (Schultz) Kindb. [SHV 20504]
bifrons (De Not.) Delg. [JSS 13-81; KLL 1393]
rigida (Hedw.) Limpr. [KLL 1379B]
Barbula
convoluta Hedw. [KLL 1367a]
Crossidium
seriatum Crum & Steere [KLL 1395, 1400]
Didymodon
australasiae (Hook. & Grev.) Zand. [JSS 13-56; KLL 1399]
brachyphyllus (Sull.) Zand. [JSS 13-31; KLL 1370]
nicholsonii Culm. [KLL 1375]
tophaceus (Brid.) Lisa [SHV 20494]
vinealis (Brid.) Zand. [JSS 13-14; KLL 1368; SHV 20501, 20503]
Eucladium
verticillatum (Brid.) B.S.G [JSS 13-55; KLL 1378; SHV 20510]
Gymnostomum
aeruginosum Sm. [JSS 010345; SHV 20505]
Microbryum
davallianum (Sm.) Zand. [JSS 13-67; SHV 20512]
starkeanum (Hedw.) Zander [KLL 1390]
Phascum
cuspidatum Hedw. [KLL 1401]
Pseudocrossidium
replicatum (Tayl.) Zand. [JSS 13-29]
Stegonia
hyalinotrichum (Card. & Thér.) Zand. [KLL 1394]
Syntrichia
bolanderi (Lesq. & James) Zand. [JSS 13-24]
inermis (Brid.) Bruch [SHV 20514]
princeps (DeNot.) Mitt. [JSS 13-25; SHV 20495, 20497]
ruralis (Hedw.) Web. & Mohr [JSS 13-74]
Timmiella
anomala (BSG) Limpr. [JSS 13-10, 13-46; SHV 20490, 20492, 20498, 20508]
Tortula
atrovirens (Sm.) Lindb. [JSS 13-13; KLL 1374]
brevipes (Lesq.) Broth.
[JSS 13-48, 13-76; KLL 1365, 1385, 1396 1402; SHV 20502, 20511]
californica Bartr. [KLL 1392]
obtusifolia (Schwaegr.) Math. [KLL
1376]
protobryoides Zand. [KLL 1379a]
Weissia
andrewsii Bartr. [JSS 13-63]
Rhabdoweisiaceae
Dichodontium
pellucidum (Hedw.) Schimp. [KLL 1369,
1386]
Discusion
Sixty-five
species of mosses in thirty-two genera and twelve families were collected
and identified from Santa Margarita Ecological Reserve. Twelve liverwort taxa
were found, representing seven genera and six families, and two hornwort taxa
were found. These collections represents over 10%
of the moss flora known from California. The liverworts,
though not as diverse in the region, are represented by several distinct taxa.
Complex thalloid taxa are especially well represented. Eleven mosses, representing about 17% of the total
number found, are here documented for the first time in Southern California
(Bryum pyriferum, Eurynchium hians, Scleropodium colpophyllum, Pleuridium acuminatum,
Fissidens sublimbatus, Grimmia lisae, Didymodon australasiae, Pseudocrossidium
replicatum, Schizymenium shevockii, Tortula obtusifolia,
and Weissia andrewsii). Another three
species in Bryum (sens. lat.) are new to science. Two liverwort species are rare narrow endemic taxa (Sphaerocarpus drewei, Geothallus tuberosus),
and two species of mosses are considered rare in the area but have wider
geographic distributions (Stegonia hyalinotrichum, Homalothecium arenarium).
The bryophyte flora of the area is a composite of several floristically
distinct bioregions that overlap to produce a relatively high diversity
for such an arid environment. First, the moss flora of the Santa Margarita Ecological
Reserve region exhibits an affinity to the Mojave Desert, which occurs to
the north and east. In excess of a third (36%) of the
53 species reported also occur in the Mojave (Harthill et al. 1979; Stark
& Whittemore 2000). The majority (10) of these
are in the families Pottiaceae and Grimmiaceae, each with xeric tendencies. However, this trend is absent among the liverworts, with
only two species in common with the Mojave Desert, Targionia hypophylla
and Asterella californica.
The relatively high liverwort diversity reported here probably results
from the cooler, wetter winter climate along with significantly greater precipitation
in this region compared to the deserts. The latter
species, although reported from the Mojave, may actually represent an undescribed
species of Targionia endemic to the Mojave Desert (Whittemore
1996). Given the proximity of the Santa Margarita
Reserve to the Mojave Desert, it will thus be of interest to compare the
Santa Margarita specimens with those to the north and east, near Joshua Tree
National Park.
Crossidium seriatum had previously been reported from southern California
from only a single locality, in a desert region of eastern San Diego County. This species is a noted gypsophile and globally rare species, known from fewer than 10 populations worldwide
(Baja California, Anza‑Borrego Desert, southern Nevada, southern Arizona,
and Spain; Zander 1977, Cano et al. 1992, Stark & Whittemore 1992, Stark
2000). Finding this species outside of its normal gypsum
habitat is notable.
Secondly, a pacific coastal element is also strongly represented in the
Santa Margarita Ecological Reserve bryoflora. That biogeographic element
includes Anacolia menziesii, Syntrichia princeps, Homalthecium
arenarium, and Claopodium whippleanum, all of which are broadly
distributed along the Pacific coast. In addition to these discrete biogeographic
elements, several cosmopolitan species are, not surprisingly, found in the
reserve. These include Funaria hygrometrica, Anomobryum argenteum
/ A. lanatum, Rosulabryum capillare, Ceratodon purpureus,
Grimmia pulvinata, Didymodon vinealis, and Syntrichia ruralis.
The Bryaceae is a particularly well represented family in the area, with
thirteen species in three currently recognized genera. Bryum caespiticium, B.bicolor, B.
pseudotriquetrum, B. gemmiparum are all relatively frequent
in the area, and their collection comes as no surprise. Bryum gemmiluscens was a surprise because it was the most common
species at the reserve and is relatively less frequent in surrounding areas. It is apparently under-collected in California and will
likely be found as a prominent component of the bryoflora in other areas as
well. Bryum pyriferum was a real surprise. It has only recently been found for the first time in
California, in the San Francisco Bay area (Shevock & Toren, 2001), and
the Santa Margarita collection is only the fifth known location for this
species in North America.
Three undescribed species of Bryum (sens. lat.)were found in addition to
the known taxa. "Species B" is an interesting little
species that is related to the Bryum bicolor
group. To date it is known only from an area in west
central California and from the Santa Margarita Ecological Reserve. It is very likely a California endemic.
"Species C" and "Species D" are distinctive undescribed species found
in rock crevices. "Species D" is somewhat similar to
Brachymenium exile, a pantropical species that is known from
Mexico. As far as "species D" is known, it is also a California endemic.
In addition to these undescribed species, other collections provided identification
challenges and could not be readily placed into any know taxon. One collection in particular (Kellman 1394) proved very
reticent to diagnosis. Superficially similar to Physcomitrella,
closer inspection demonstrated that it could not be a member of Funariaceae.
Morphological details were assessed by Bruce Allen as follows (personal communication). Comal leaves are broadly obovate and concave, approximately
2 mm (including the pilifer), entire, and bruptly tapered
to a slender and essentially hyaline pilifer (which is about 0.3x the
total leaf length). The upper laminal cells are thin-walled,
smooth, and certainly could pass as Funariaceae. The
costa is excurrent into the pilifer. There is a definite
swelling on the adaxial surface below the base of the pilifer which is suggestive
of rudimentary adaxial lamellae such as those found in certain members of
Pottiaceae, such as Crossidium. The calpytra is mitrate,
covering only the apex of the capsule, and the capsule is broadly ellipsoid,
bluntly beaked at the apex, about 0.8 mm long, and inoperculate, with no
suggestion of a line of dehiscence. The exothecial
cells very thin-walled, quadrate to five-sided and approximately equal in
length and width, about 30-45 m in the greater dimension. Stomata are few, restricted
to capsule base, definitely 2-celled, and superficial. Immature
spores are essentially hyaline, about 15-18 m in diameter, and without surface ornamentation,
which would likely be obvious if the specimen were a Physcomitrella.
The thin-walled inoperculate capsule, mitrate calyptra, and the laminal
areolation are highly suggestive of Funariaceae. However,
the definite swelling on the adaxial surface of the costa (just below the
base of the pilifer, and suggestive of rudimentary adaxial lamellae), and
especially the 2-celled stomata indicate this material does not belong to
that family (Allen, personal communication). The final
determination is that this specimen represents Stegonia,
a relatively rare member of the Pottiaceae in this region.
It is apparent from the results of this four-day intensive field meet that
the bryophyte flora in the west is still far from well understood and documented. One of the objectives of this annual meeting is to contribute
to exploration and description of the flora. Founded
in 1996, SO BE FREE is a series of West Coast forays started by the Bryolab
at UC Berkeley, but open to all botanists. The main
focus is on bryophytes, but we also encourage experts on macroalgae, mushrooms,
lichens, ferns, and flowering plants to come along. We
welcome specialists as well as generalists, or amateurs who are interested
in an overview. It is held each spring, associated
with Spring Break at universities. This distinguishes
it from the eastern bryological forays (the Andrews and the Blomquist Forays)
which are held in the fall, and allows the occasional easterner,
desperate as they often are to see plants and blue sky in the spring, a chance
to attend.
The usual tradition is to have a four-day, three-night schedule with communal
cooking and eating in inexpensive and remote biological field stations. Evening slide shows and informal talks are presented as
well as keying sessions with microscopes. In addition
to seeing interesting wild areas and learning new plants,
important
goals for SO BE FREE include keeping west coast bryologists (and friends)
in touch with each other and teaching and encouraging students and amateurs.
For a
glimpse of some of the past six outings, consult the SO BE FREE web site:
http://ucjeps.herb.berkeley.edu/bryolab/trips/sobefree.html
In 2002 the Seventh Annual SO BE FREE will meet in the foothills of the
Mayacamas Mountains at the Hopland Research and Extension Center (HREC; see
http://danrrec.ucdavis.edu/hopland/home_page.html). HREC is the University
of California's principal field research facility for agriculture and natural
resources in the North Coast region. The Center extends
over more than 5,300 acres. A diversity of soils,
plant and animal communities, and elevations makes HREC representative of
many parts of the Coast Range in northwestern California.
The meeting will convene Saturday, 23 March 2002 at the Hopland Research
and Extension Center at 4:00pm. Detailed directions
about how to get there will be sent out to registrants. One
full day we will be spent exploring the Mendocino coast, seeing redwood
forest, deep canyons, and the famous Mendocino pygmy forest. The other full
day we will go to drier interior habitats in Lake County.
In the evenings we will have slide shows about bryophytes and lichens,
and current research efforts of the participants followed by identification
sessions. Please inform Brent Mishler of your firm intent to attend the foray,
by February 1st, 2002, at: bmishler@socrates.berkeley.edu, or by telephone 510-642-6810.
Please include your mailing address and phone number, and note any
special restrictions you might have on diet.
Whenever and wherever bryologists turn out in force to concentrate and coordinate
their observations of the flora, new taxa, new distribution records, and rare
and little-known organisms come to light. This is
clearly demonstrated by the results of the 2001 meeting of SO BE FREE, and
we anticipate that the 2002 meeting will just as productive. All are invited.
Literature Cited
Anderson,
L. E., H. A. Crum & W. R. Buck. 2000. List of mosses of North America
north of Mexico. The Bryologist 93(4): 448-499.
Bowers,
f. D., C. Delgadillo M., A. J. Sharp 1976. The mosses of Baja California.
J. Hattori Bot. Lab. 40: 397-410.
Buck,
W. R. & B. Goffinet. 2000. Morphology and classification of mosses. Chapter3:
72-123. In Shaw & Goffinet (eds.). Bryophyte Biology. Cambridge Univ.
Press.
Cano,
M. J., J. Guerra, and R. M. Ros. 1992. Crossidium seriatum (Pottiaceae,
Musci) new to Europe. Bryologist 95: 280‑282.
Harthill,
M.P., D.M. Long, and B.D. Mishler. 1979. Preliminary list of southern California
mosses. Bryologist 82(2): 260-267.
Sharp,
A. J., H. Crum, and P. M. Eckel (eds.) 1994. The Moss Flora of Mexico. Memoirs
of the New York Botanical Garden, Vol. 69.
Shevock,
J. R. and D. Toren 2001. A specimen-based catalogue of mosses for the city
and county of San Francisco, California. Madroño 48(1): 1-16.
Stark,
L. R. 2000. Bryophyte inventory of the North Shore region of Lake Mead, Clark
County, Nevada. Report, National Park Service and the U.S. Bureau of Land
Management.
Stark,
L. R. and A. T. Whittemore. 2000. Bryophytes from the northern Mojave Desert.
Southwestern Naturalist 45:226-232.
Stark,
L. R. and A. T. Whittemore. 1992. Additions to the Bryoflora of Southern California.
Bryologist 95(1): 65-67.
Whittemore, A. T. 1996. The taxonom of Targionia (Targioniaceae)
in North America (abstract). American Journal of Botany 83(6 suppl.) 22-23.
Zander,
R. H. 1993. Genera of the Pottiacea: Mosses of harsh environments. Bull. Buff.
Soc. of Nat. Sci. Vol. 32. 378 pp. New York.
Zander,
R. H. 1977. Crossidium seriatum found in the U.S.A.
Bryologist 80: 1709‑171.
Acknowledgements We would like to thank Bruce Allen for his insightful comments
on KLL 1394, Dan Norris for expert assistance and concurrence on several key
determinations, Brent Mishler for coordinating the Sixth Annual SOBEFREE,
and Claudia Luke for graciously hosting our meeting.