Grimmia vaginulata, (Bryopsida, Grimmiaceae) a New Species from the Central Coast of California

2011·7.7 MB·English

by  Kenneth Kellman

Madrono, Vol. 58, No. 3, pp. 190-198, 2011
GRIMMIA VAGINULATA, (BRYOPSIDA, GRIMMIACEAE) A NEW SPECIES
FROM THE CENTRAL COAST OF CALIFORNIA
Kenneth Kellman
California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118
[email protected]
Abstract
A new California endemic species, Grimmia vaginulata, is described and illustrated. It is
characterized by its very small size, julaceous habit, immersed capsule on a straight, centrally
inserted seta, verylarge annulus, keeled unistratose leaves, autoicous sexuality, and a largecylindrical
ochrea atop the vaginula that sheathes the seta to the base of the capsule. The differentiation and
ecology ofthe new species is discussed.
Key Words: California, endemic moss, Grimmia vaginulata, Grimmiaceae, new species, ochrea,
vaginula.
In the summer of 2008, prior to the annual the northern intersection with SR 236, elev.
meeting of the American Bryological and Li- 685 m, 37T2'40"N, 122°12'42"W, 13 Aug2008.
chenological Society, Dale Vitt asked to see Kellman, Vitt, & Shevock5869(holotype: CAS).
&OrtGhooftfriicnleitwninketUhieuafnieilid.D.DurHi.ngNotrhraits,shSohretvotrcikp sedSpdeicffieerst aGbrismtmatiuaracbaprielvliaotrae,DfeoHiNsotsatratiusmaafdfiinidso,
Dale noticed tiny little plants growing on the vel sicco persistenter julaceis, ochrea cylindracea
same sandstone boulders that supported the longiore usque ad prope basem capsulae exten-
Orthotrichum. These plants were so scattered
denti differt.
and insignificant that they appeared to be Plants up to 2 mm tall, scattered to veryloosely
iminature stems of a small Grimmia, and not
tufted (Fig. 5), simple, or with a short, tightly
worth collecting. That impression proved incor- appressed branch; green when young, tan or
rect. Upon microscopic examination, it was white in age; julaceous wet or dry. Lower and
immediately obvious that these plants represent-
perigonial leaves short ovate, 1:1, muticous,
ed a remarkable plant that was unlike any other increasing in length upwards. Upper stem and
Grimmia in North America. perichaetial leaves (Fig. 2B) obovate or eUiptical,
Chief among the distinguishing characteristics soft when moist, weakly keeled concave, uni-
was a transparent extension of the vaginula that stratose throughout, with at least some portion of i
extended to the base of the capsule. This the leafmargin hyaline, most common at the base |
extension, known as an ochrea, is not present in and often extending to midleaf or slightly [
any North American Grimmia, and it is not beyond; 1-1.65 mm long X 0.4—0.75 mm wide
mentioned in any North and Central American (without the awn); the keel is even less pro-
Grimmia literature (Flowers 1973; Crum and nounced in transverse section, appearing more
Anderson 1981; Crum 1994; Hastings and convex, even tubular, especially on non-sporo-
G20r0e2v)e.nTo200t7h;e cMausnuaolzst1u9d9e9n;t Moufhtohze gaenndus,Altlheins pslhiygthitclyplaacnutmsin(Faitge..4)B;asaaplexjuaxctuatceosotralocccealslsionsahlolryt
novel character suggested the possibility ofa new rectangular with thin to moderately thick,
genus. That hypothesis was supported by the straight; relatively uniform across the base, but
scattered growth habit. A wider examination of outer basal cells often somewhat narrower in 1 to
Grimmia was undertaken resulting in the expan- 3 marginal rows. Distal and medial cells decreas- •
sion of the North American parameters of the ing in length gradually from the base to the apex,
genus. 1-1.5(2.5):1, 12-16(26) |am long X 7-13 )im wide;
rectangular, triangular, or irregularly polygonal,
i
Taxonomy thick walled, often with some portion of the
lumen rounded creating small trigones, not or
Grimmia vaginulata Kellman, sp. nov. (Figs. lA, weakly sinuose; in transverse section plane to
IC, 2, 3, 4, 5) Type: USA, Cahfornia, Santa slightly bulging. Margins weakly recurved on one
Cruz Co., Big Basin Redwoods State Park, on orboth sides; unistratose. Costanarrow, to40|Lim
!
dry vertical walls of calcareous sandstone wide at the base, broadening toward the apex;
bouldeis eroded from the sandstone bedrock excurrent in a hyaline, weakly toothed awn ;
i
in open chaparral, above the Basin Trail and shorter than the lamina, decurrent at most 1-2
i
below China Grade Road ca. 1.5 mi. beyond cells down the margin, and those often projecting
2011] KELLMAN: A NEW SPECIES OF GRIMMIA 191
Fig. 1. A. Grimmia vagmulatawithleaves strippedexposing the sporophyte. (O) ochrea (V)entirevaginula. Scale
bar = 500|Lim(fromthetype); B. G. capillatawithleaves strippedexposingthe sporophyte. (O) ochrea. Scale bar =
500 \xm {Hcmdel-Mazetti J778, FH: syntype of G. niesopotamica); C. Detail of typical upper leaf apex of G.
vaginulata Scale bar = 80 |im; D. Detail oftypical upper leafapex of G. ccipillata Scale bar = 175 |im.
mm
as short, blunt teeth; costa in transverse section at measured with ochrea, 0.5-0.6 long, epider-
midleaf with two homogenous rows of cells, 2 mal cells irregular, about 2-3:1, with very thin,
cells wide adaxially. Gonioautoicous or cladau- straight walls; distally with a cylindrical ochrea
toicous. Calyptra irregularly crenate at the base, surrounding but not connate to the seta and
conical or campanulate, hyaline at the extreme flaring just below the base of the capsule, 340-
apex and at the base, naked and smooth, just 430 |am long. Seta straight, attached to the center
mm
covering the operculum. Vaginula (Fig. lA), of the capsule, 0.5 long. Capsule immersed
MADRONO
192 [Vol. 58
G. vaginulata
Fig. 2. A. Sporophytic plants of Grimmia vaginulata and G. capillata showing size difference. Marks are
apart; B. G. vaginulata, two upper and perichaetial leaves. Scale bar = 500 |j.m.
with only the operculum exposed, irregularly Grimmia vaginulata resembles G. anodon Bruch
wrinkled when dry, slightly wrinkled when wet, & Schimp. and G. plagiopodia Hedw. in leaf
obloid, 0.9 mm long X 0.6 mm wide, abruptly shape. However, G. vaginulata remainsjulaceous
contracted to the seta. Annulus (Fig. 3) of3 rows wet or dry, and is a much smaller and narrower
of differentiated, transparent, thick-walled en- plant. The straight, centrally inserted seta of G.
larged and elongated cells; remaining on the urn vaginulata contrasts with the sigmoid, eccentri-
after dehiscence, but gradually falling off in cally inserted seta that characterizes both G.
sections. Operculum mammilose to low conical, anodon and G. plagiopodia. The extremely dis-
crenulate to erose at the base. Exothecial cells sected peristome resembles G. orbicularis Bruch
irregularly rectangular to hexagonal, thin-walled, and G. moxleyi R.S. Williams. Grimmia orbicu-
almost transparent when mature, easily revealing laris shares the unistratose margins, and the
the stalked theca within. Stomata present. Peri- autoicous sexuality with G. vaginulata, but the
stome (Fig. 3) of 16 orange-red cribrose-dissected awns of G. orbicularis are evenly distributed
teeth, irregularly divided nearly to the base into along the stem. Grimmia moxleyi has awns
3^
strongly spiculose filaments, ca. 185 |j,m long. restricted to the upper leaves like G. vaginulata,
Spores smooth, 10-13 |im in diameter. but the leafmargins are bistratose. Furthermore,
The specific epithet refers to the persistent both G. orbicularis and G. moxleyi have exserted
cylindrical ochrea atop the vaginula, which capsules on arcuate setae. Again, both are much
extends nearly to the base ofthe capsule. larger than G. vagiimlata.
Bothgametophytically, and sporophytically, G.
Identification and vaginulata appears most closely related to G.
Taxonomic Relationships capillata De Not., a species scattered around the
Mediterranean Sea (Greven2003). They sharethe
Grimmia vaginulata can be distinguished from keeled, entirely unistratose lamina with the
all other North American Grimmia by the margins at least somewhat recurved on one or
following combination of characters: very small two sides, a costa that broadens in the distal half
plants that arejulaceous wet or dry, upper leaves of the leaf, perichaetial leaves with proximal
with hyaline margins, dissected peristome, and a hyaline areas, and autoicous sexuality. Sporophy-
conspicuous and persistent ochrea sheathing the tically, the two taxa are very close. Lastly, both
entire seta to just below the base ofthe capsule. prefer calcareous substrates. Again, G. vaginulata
2011] KELLMAN: A NEW SPECIES OF GRIMMIA 193
0
Fig. 3. Detail ofannulus and peristome of Grimmia vaginulata. Scale bar = 120 |im.
isamuchsmallerplantthatstaystightlyjulaceous
My
wet ordry. examination of16 specimens ofG.
capillata and G. mesopotamica Schiffn. (a syno-
nym of G. capillata fide Munoz and Pando 2000;
Greven 2003) from FU, MUB, and NY show
plant size in G. capillata averages around 7 mm,
dwarfing G. vaginulata. (Fig. 2A) The dry leaves
of G. capillata are erect to erect-patent and are
usually strongly keeled. Thus the costae project
from the rest ofthe leaves giving the dry plants a
more tumid and textured appearance. When
moist, the leaves of G. capillata are at least
erect-patent, and are relatively easy to dissect
from the stem. The tiny and tightly appressed
leaves of G. vaginulata are very difficult to strip
for examination without tearing. Leaf size is
another point ofseparation, with the upper stem
mm
leaves of G. capillata measuring 2-3 long
(Cortini Pedrotti 2001; Ignatova and Munoz
2004), while the much smaller G. vaginulata has
leaves with a maximum length of 1.65 mm. In G.
capillata, themore orless rounded apex (Fig. ID)
is the most commonly hyahne portion ofthe leaf,
while in G. vaginulata, the margins are hyaline, Fig. 4. Drawings of leaf cross sections, Grimmia
and the apex is acute. There is some small portion vaginulata: a) leaffrom sterile plant taken at midleaf; b,
at the base ofthe awn that is hyaline, but at most c, d) upper leaf from sporophytic plant. Scale bar =
it is one to two cells down the margin (Fig. IC). 100 |im.
MADRONO
194 [Vol. 58
Fig. 5. Grimmia vaginiilata. Photograph ofgrowth habit on sandstone.
Although evidence is admittedly sparse, with the vaginulata. G. pseudoanodon has no peristome, its
onlyknownpopulationbeingthetype, thegrowth cauline leaves are not reduced down the stem,
habit of G. vagimdata adds another basis for beinglinear to lanceolate and bearingawns to the
separation. In G. vaginulata, the plants are at best base of the plant. They are patent to spreading
loosely associated (Fig. 5), not forming a contin- when moist, and are easy to separate from the
uous turf or clump. In G. capillata, the plants stem. Similar to G. capillata, the apex ofthe leaf
form dense green mats of sterile plants inter- is hyaline for 15-20 cells, and the awn is often
spersed with a few fertile plants (Cortini Pedrotti decurrent down the margins of the leaf. Most
2001; Greven 2003; Heyn and Herrnstadt 2004; leaves of G. pseudoanodon show bistratose
Ignatova and Muiioz 2004). The base of the margins. The vaginula of G. pseudoanodon is
capsule in G. capillata is less abruptly contracted extended, demonstrated by the aborted archego-
to the seta than is seen in G. vaginulata. Lastly, in nia scattered throughout its length, however, the
G. capillatatheochrea,whilepronounced,extends ochrea is at most represented by a few hyaline
only halfway up the seta, not to the base of the flaps at the apex of the vaginula, far from the
capsule as in G. vaginulata. (Figs. lA, B) tube sheathing the seta of G. vaginulata. Inter-
Grimmia pseudoanoclon Deguchi, described in estingly, slight pressure expels the seta from the
1987 from Peru (Deguchi 1987), is another small end ofthe vaginula, leaving a conical stub at the
autoicous plant (ca 5 mm tall) with immersed base of the seta.
sporophytes with a centrally attached, straight
seta, and like G. vaginulata, the base ofits capsule Generic Taxonomy
is abruptly contracted. Examination ofan isotype
from NY, showed several important differences It is clear that this new plant belongs in the
from G. vaginulata. First, G. pseudoanoclon grows Grimmiaceae. The only other familial possibility
in cushions, contrasting the scattered habit of G. is Ptychomitriaceae, and that is ruled out by the
.
2011] KELLMAN: A NEW SPECIES OF GRIMMIA 195
unlobed calyptra. Within Grimmiaceae, other wrote "I believe the ochrea is a fairly common
genera can easily be dismissed as well. The large feature of the neotropical species of Grimmia. It
annulus, and the operculum falling independent can be demonstrated in G. americana, G.
ofthe columnella deny placement in Schistidium. donniana and G. elongata. It is particularly
Although the cribrose peristome resembles those evident in G. anodon."" But more convincing
of both Coscinodon and JaffueJiobryum, and the examples can be found in the persistent ochreae
autoicous sexuality, unistratose lamina and large of G. capillata and G. involucrata Cardot.
annulus further support placement in Jaffuelio- Being clear then that this new plant properly
bryum, both genera require a pleated and belongs in Grimmia, it is necessary to at least
somewhat sheathing calyptra (Churchill 1987; explore its relationships within the genus. The
Hastings 2007; Spence 2007). The calyptra of G. historyofthegenus Grimmia, s.h is a tortured and
vaginulata is so short that it does not even split or complex story of confused concepts, alternating
tear during capsule development, sitting instead periods of splitting and lumping, and regional
just atop the operculum. Racomitrium {sensu lato) authors describing numerous plants that have
is ruled out by the lack of sinuosity in the cell been later synonymized in worldwide treatments
walls of the lamina or the vaginula (Deguchi (Munoz and Pando 2000; Greven 2003). Deguchi
1978). (1978) and Ochyra et al. (2003) both provide a
Not only can we suggest placement in Grimmia good summary of this nomenclaturally difficult
by elimination, but the dramatic characters subject. Although Hernandez-Maqueda et al.
displayed by G. vaginulata fit well within the (2008) casts some doubt on Coscinodon, most
limits ofvariation ofplants already in that genus. modern authorities agree on the separation of
Deguchi (1978) describes the "Affinis type" Jaffueliobryum, Schistidium, and Coscinodon from
annulus as "well differentiated, composed of (2) Grimmia, but there is disagreement whether
3^ rows of cells, which are thick walled, but Grimmia itself requires further subdivision.
transparent, and becoming increasingly larger Ochyra et al. (2003) proposed to split Grimmia
from the lower to the upper rows. ...Upper rows into fivegenera, witheachgenuscorrespondingin
ofcells ofthe annulus are also removed when the concept with the subgenera reluctantly suggested
lid falls, but their lower rows usually remain by Hastings and Greven (2007). Both systems are
attached to the orifice ofthe urn, disjoining little rooted in the work of Hagen (1909) and are
by little in the course of time." This is a perfect summarized in Table 1
description of the annulus in G. vaginulata. He Grimmia vaginulata can be ruled out of
attributes this "Affinis type" annulus to "G. Gasterogrimmia based on the straight, centrally
affinis, G. anomala, G. apiculata, G. brachydic- attached seta. Litoneuron requires 2-stratose,
tyon, G. curvata, G. olympica, and G. pilifera."" concave leaves with the costa not prominently
Munoz (1999) assigns this annulus type to "G. projecting abaxially: Grimmia vaginulata meets
involucrata, G. longirostris, G. poecilostoma, and none of these criteria. An expanded Guembelia
G. trichophylla'' in the introductory section, and that included unistratose leaves would fit all
14 other taxa in the species descriptions. characters of G. vaginulata, but if one were to
The ochrea is a structure that is poorly accept the Ochyra et al. (2003) concept of
understood and has received little taxonomic Dryptodon, which includes plants with short
discussion. Deguchi (1978) makes it clear that the straight setae and phcate or wrinkled capsules,
ochrea is only a section of the vaginula, but then G. vaginulata would belong there. In fact,
Magill (1990) defines ochrea as "vaginula, or Ochyra et al. (2003) places G. capillata in
upper part of vaginula". The haploid vaginula Dryptodon.
includes cells originating from both the archego- The difficulty is compounded, however, by
nium and the upper part of the stem (Deguchi statements by various authors suggesting that
1978). Deguchi's proof of this contention is the pairs of plants within Grimmia s.l. are closely
"occurrence of aborted archegonia at the basal related, but subsequent authors treat each
part ofthe vaginule" (Fig. lA). member of the pair in separate subgenera or
Maier (2002) discusses the vaginula and the genera. For example, G. capillata is often paired
ochrea and describes Grimmia donniana Sm. ex with G. crinita Brid. (Greven 2003; Ignatova and
Spruce with "ochrea broad", with most other Munoz2004) and in fact DeNotaris (1836, 1838),
species with ochrea "short" or "small". Some who first described G. capillata in 1836, reduced it
species such as G. orbicularis (a close relative of to a variety of G. crinita in 1838, and Maier
the aforementioned G. moxleyi) she describes (2010) reduced G. capillata into synonymy with
with "vaginula 0.8 mm, with ochrea." Unfortu- G. crinita. In this pair, G. crinita is universally
nately, she does not provide illustrations to placed in Gasterogrimmia {Grimmia sensu Ochyra
inform that aspect of her excellent descriptions, et al. 2003) but as stated above G. capillata is
nor does she describe how she measures the placed in Dryptodon. These cross subgeneric or
vaginula. Delgadillo (Universidad Nacional generic affinities, as well as the taxa that defy
Autonoma de Mexico, personal communication) placement within any system, illustrate the
MADRONO
196 [Vol. 58
Table 1. Comparisonof Taxonomic Treatmentsof Grimmia.
Hastings and Greven (2007) Ochyra et al. (2003) Description
Gnsterogrifiimia Gritntnici Capsules immersed, smooth, seta
sigmoid and eccentrically attached
tothecapsule,leaves 1-or2-stratose
that are concave or concave-keeled.
Guembelia Orthogrimmia Capsules immersed to exserted,
smooth, seta straight and centrally
attached to the capsule, leaves
mostly 2-stratose and keeled.
Litoneuron Guembelia Capsules exserted or emergent,
smooth, seta straight and centrally
attached to the capsule, leaves 2-
stratose that are concave with the
costa not prominent dorsally.
Rhabdogrimmia Dryptodon Capsules emergent to exserted, ribbed,
seta arcuate and centrally attached
to the capsule, leaves mostly 1-
stratose (2-stratose at the margins)
and keeled.
complexity ofGrimmias.l. It is likely that this will Adenostoma fasciculatum Hook. & Arn., Arcto-
only be sorted out by a massive study combining staphylos tomentosa (Pursh.) Lindl., Ceanothus
worldwide genetic and morphological data. cuneatus (Hook.) Nutt., C. papillosus Torr. & A.
The very dramatic and persistent ochrea in G. Gray, and Eriodictyon californicum Hook. &
vaginulata opens the question of the importance Arn.. The climate is Mediterranean with moder-
ofthe ochrea in Grimmia taxonomy. Only Maier ate but rainy winters, and hot rainless summers
(2002) and Deguchi (1978) discuss this character. (Kellman 2003). The elevation is sHghtly below
Deguchi briefly mentions the ochrea in his 700 meters, placing the site above all but the
introduction, and then does not use it in his thickest summer maritime fog. The lack of
species descriptions. It is ignored in Crum (1994), summer fog keeps these sites very hot and dry
Muiioz (1999), Cortini Pedrotti (2001), Greven until the winter storms arrive.
(2003) Ignatovaand Munoz(2004), and Hastings One of the more peculiar aspects of this very
,
and Greven (2007). Maier (2002) lists G. donniana interesting plant is the substrate. With the
with ochrea ''broad", and G. elatior with ochrea exception of one recent collection from marble
"distinct". Many other species were described rock {Kellman, Shevock& Lodder6133 [CAS]) the
with ochrea "small" or the character was ignored boulders are also the sole known substrate of
A
altogether. quickexaminationoffivespecimens Orthotrichum kellmanii, another rare coastal
in CAS annotated by R. Hastings as G. domiiana Cahfornia endemic (Norris et al. 2004). The
displayed three with no ochrea, one veryjuvenile bedrock was deposited in the Upper Eocene, and
capsulewithanochreaalreadydisintegrating, and is part of the Butano Sandstone Formation
one specimen with a persistent ochrea about as (Brabb 1989), but the geologic history of the
longas broad at the base ofa mature sporophyte. boulders is not known. Presumably, the rocks
Thus,itseemsthat thischaracterbyitselfdoesnot were formed and eroded into their rounded shape
seem to be rehable enough to be useful for some time prior to the Eocene deposits that
separating G. donniana, one of the few taxa that formed the bedrock. In Santa Cruz Co., the
Maierthoughthadadistinctiveochrea; itislikely, Butano Formation is exposed along the western
therefore, that the character is not useful beyond slopes of the highest mountains on the eastern
the G. capillata-vaginulata group. Nonetheless, side ofthe county (Brabb 1989), but the boulders
further study of such taxa as G. involucrata may are only present in a small patch in Big Basin
show that the ochrea could be an additional Redwoods State Park. There is also a small field
character useful for identification. ofthe boulders in southern San Mateo Co., also
in Butano State Park. Recently, this same
Ecology and Distribution formation with exposed boulders has been
found in Monterey Co. within the Ventana
Grimmia vaginulata has thus far only been Wilderness of the Los Padres National Forest.
found on vertical or underhanging surfaces on Orthotrichum kellmanii was collected here al-
calcareous sandstone boulders that have eroded though much ofthe area remains to be surveyed.
out of less calcareous sandstone bedrock of the The new population greatly expanded its range
Butano Formation. These rocks are scattered to the south. Although access to these sandstone
through a variably dense chaparral comprised of outcrops was difficult due to hiking through
2011] KELLMAN: A NEW SPECIES OF GRIMMIA 197
dense stands ofchaparral, the area subsequently full description. Of course, it is always possible
burned in the summer of 2008. This area is now that further collections from new locations could
among the highest priority sites to survey for G. alter the concept of G. vaginulata. Of all the
vaginu/ata-and to document the extent of Ortho- characters discussed above, the most likely
trichum keUmanii in this area. features to change would be the size and the
Associated bryophytes growing on and around growth form. It is possible that other populations
the boulders include Amphidium californicum of G.vaginulata could be composed of larger
(Hampe ex Miill. Hal.) Broth., Antitrichia cali- plants. Arguing against that possibility is the fact
fornica Sull. in Lesq., Gemmabryum californicum that both sterile and fruiting plants exhibit the
(Sull.) Spence, Didymodon vinealis (Brid.) R.H. samesizeandgrowth form in thetypepopulation.
Zander, Tortula niuralis Hedw., Grimmia torenii It seems more likely that G. vaginulata could be
Hastings, G. pulvinata (Hedw.) J. E. Smith, found growing in turfs or small cushions.
Gymnostomum calcareum Nees & Hornsch., G. However, turf and cushion formation require
viridulum Brid., Orthotrichum kellmanii, and the survival ofthe originating stems. There are no
Cephaloziella divaricata (Sm.) Warnst. subapical innovations in the type population
which suggest that this species may be short lived
Conservation and that individualsdieaftersexual reproduction.
Acknowledgments
At the type location, G. vaginulata has been
found on only four out of several hundred I am indebted to Jim Shevock and Dan Norris for
boulders, and these fit within a circle of ten teachingmehowtodo the research necessary towritea
meters. It is imperative that no furthercollections paper of this type. They may or may not have been
should be made until more populations are awarethat theywereteachingmethesemethods, butan
found. Additional surveys need to be conducted interested student learns as much by observation as by
throughout the highly restricted habitat within laelcstoulriekeatnodtdhiasnckusRsiooxna.nInehoHnaosrtitnhgesirfoerxatemapclhei.ngImweoutlhde
the Butano Formation within State Park Lands basic understanding of Grimmia, and for a thorough
and a federally designated wilderness area. Even reviewofthispaperresultinginmanypositivechanges. I
ifadditional occurrences are discovered, Grimmia thankHenk Grevenforhisreviewofthispaper, andthe
vaginulata will almost certainly remain a very suggestion to compare Grimmia vaginulata with G.
narrowly restricted endemic that could be ad- pseudoanodon. DavidToren deservescredit forpointing
versely impacted by stochastic events. out the illustration of Grimmia mesopotamica in the
Even though the chaparral is a fire adapted sBirmyiloaprhityitees oFfloGr.avaogfinIuslraatealatnhdatG.puctapimlleata.onTthoantkhse
ecosystem (Schoenherr 1992), the extreme rarity also to Eva Maier, and Claudio Delgadillo, for their
ofG. vaginulata leaves open the possibility offire insights on the vaginula and ochrea. I also appreciate
killing the entire known population. In March of Claudio Delgadillo and Jesus Munoz for their helpful
2009, the author visited the Bonny Doon Ecolog- comments on an earlier version of this paper. Steve
ical Preserve in Santa Cruz Co. Nine months Lodder was invaluable in providing the beautiful
before, in June of 2008, a very hot fire raced photographs. I thank Patricia Eckel for providing the
through the preserve, where sandstone rocks are Latin description. I would like to thank the curators of
scattered through a chaparral very similar to that MUB, FH, and NY for the loan of specimens of
finoupnldaceats twhheertyepethleocbartuisohnwoafsG.mivnaigminaull,atvai.rtEuavlelny pGsreiumdmoiaanocdaopnil,laatna,dGDreibmmTiraocmkesaotpCotAaSmiwcha,oaanrdraGnrgiemdmfioar
the loans.
all bryophytes were killed on the rocks, especially
those bryophytes growing on the walls and Literature Cited
underhangingsurfaces ofthe rocks. A few mosses
andliverwortssurvived onthetops ofa fewrocks. Brabb, E. E. 1989. Geologic map of Santa Cruz
Apparently the heat ofthe fire was trapped under CSeoruinest.y,U.SC.alGiefoorlnoigai.calMiSsucrevlelya,neDoeunsverI,nveCsOt.igation
the rocks and then bathed the vertical surfaces, Churchill, S. P. 1987. Systematic and biogeography
killing all plant life. It is conceivable that thick ofJaffueliobryum (Grimmiaceae). Memoirs of the
cushions, or gemmiform plants with densely New York Botanical Garden 45:691-708.
imbricate leaves could insulate at least some part CORTINI Pedrotti, C. 2001. Flora dei muschi d'ltalia:
ofa dry moss plant from the heat, but the loose Sphagnopsida, Andreaeopsida, Bryopsida (1
colonies and cylindrical plant form of G. vaginu- Parte). Antonio Delfmo Editore, Rome, Italy.
lata offers no such protection. Crum, H. 1994. Grimmiales. 386^15. in A. J. Sharp,
typTeoadnadte,inG.genveargailn,ultaataxoinsokmniocwnnovoenlltyiesfrsohmoutlhde MHe.xCircou.m,MaenmdoPi.rsM.oEfcktehle(eNdse.)w. TYhoermkosBsotflaonriacaolf
Garden 69:1-1113.
not be based on a single specimen, particularly in and L. Anderson. 1981. Mosses of Eastern
variable genera such as Grimmia. However, this North America. Columbia University Press, New
plant's combination of characters, unique to York, NY.
Grimmia in North America, along with the DeNotaris, G. 1836. Memoriedella RealeAccademia
conservation implications ofits rarity, demand a delle Scienze di Torino 39:248.
MADRONO
198 [Vol. 58
. 1838. Syllabus Muscorum in Italia et in Insulis Kellman, K. M. 2003. A catalog of the mosses of
Circumstantibus Hucusque Cognitorum, Turin, Santa Cruz County, California. Madroiio
Italy. 50:61-82.
Deguchi, H. 1978. A revision ofthe genera Griinniia, Magill, R. E. 1990. Glossarium Polyglottum Bryolo-
Schistidium and Coscinodou (Musci) of Japan. giae, a multilingual glossaryforbryology. Missouri
Journal of Science of the Hiroshima University Botanical Garden, Saint Louis, MO.
Series B, Div. 2 (Botany) 16:121-256. Maier, E. 2002. Thegenus Grinvnia (Musci, Grimmia-
. 1987. Studies on some Peruvian species ofthe ceae) in the Himalaya. Candollea 57:143-238.
Grimmiaceae. Pp. 19-74 in H. Inoue (ed.). Studies . 2010. The genus Grimmia Hedw. (Grimmia-
on cryptogams in Southern Peru. Tokai University ceae, Bryophyta) - a morphological-anatomical
Press, Tokyo, Japan. study. Boissiera 63:1-377.
Flowers, S. 1973. Mosses: Utah and the West. Munoz, J. 1999. A revision of Grimmia (Musci,
Brigham Young University Press, Provo, UT. Grimmiaceae) in the Americas. 1: Latin America.
Greven, H. C. 2003. Grimmias ofthe world. Bachuys, Annals of the Missouri Botanical Garden
Leyden, The Netherlands. 86:118-191.
Hagen, I. 1909. Forarbejder tilen Norsk lovmosflora. AND F. Pando. 2000. A world synopsis ofthe
IX. Grimmiaceae. Det Kongelige Norske Videns- genus Grimmia (Musci, Grimmiaceae). Missouri
kabers Selskabs Skrifter 1909(5):1-94. Botanical Garden Press, St. Louis, MO.
Hastings, R. I. 2007. Coscinodou. Pp. 258 262 in and B. Allen. 2002. Grimmia. Pp. 221-231 in
Flora of North America Editorial Committee B. Allen (ed.). Moss flora of Central America,
(eds.). Flora of North America North of Mexico, Part 2. Encalyptaceae-Ortliotrichaceae. Missouri
Vol. 27. Oxford University Press, New York, NY Botanical Garden Press, Saint Louis, MO.
, H. C. Greven. 2007. Grinvnia. Pp. 225 258 /// NoRRis, D. H., J. R. Shevock, and B. Goffinet.
Flora of North America Editorial Committee 2004. Orthotrichum kelJmanii (Bryopsida, Ortho-
(eds.). Flora of North America North of Mexico, trichaceae), a remarkable new species from the
Vol. 27. Oxford University Press, New York, NY. central coast of California. The Bryologist
Hernandez-Maqueda, R., D. Quandt, O. Werner, 107:209-214.
AND J. Munoz. 2008. Phylogeny and classification OCHYRA, R., J. ZARNOWIEC, AND H. BEDNAREK-
of the GrimmiaceaelPtychomitriaceae complex OCHYRA. 2003. Census catalog of Polish mosses.
{Bryophyta) inferred from cpDNA. Molecular Polish Academy ofSciences, Krakow, Poland.
Phylogenetics and Evolution 46:863-877. ScHOENHERR, A. A. 1992. A natural history of
Heyn, C. C. and I. Herrnstadt. 2004. The California. University ofCalifornia Press, Berkeley
bryophyte flora of Israel and adjacent regions. and Los Angeles, CA.
The Israel Academy of Sciences and Humanities, Spence, J. R. 2007. Jaffueliobryum. Pp. 262-264 in
Jerusalem, Israel. Flora of North America Editorial Committee
Ignatova, E. a. and J. Munoz. 2004. The genus (eds.). Flora of North America North of Mexico,
Grimmia in Russia. Arctoa 13:101-182. Vol. 27. Oxford University Press, New York, NY.