Qk M. Chapman
Vol. 30, pp. 179-184 December 1, 1917
PROCEEDINGS
OFTHE
BIOLOGICAL SOCIETY OF WASHINGTON
NOTES ON WESTERN SPECIES OF PELLAEA.*
BY WILLIAM
R.MAXON.
The
following notes relate chiefly to thetaxonomy
ofthe southwestern
ferns formany
years loosely associatedunder the
name
PellaeaWrightiana Hook.,
towhich Christensent has
recentlyassigned
thename
Pellaeamucronata D.
C.Eaton,
in error. Itisnecessary
toapply the name mucronata
toa
quite different plant,namely the
California specieslong passing
as P. ornithopusHook., and
to recognize three distinct species in the PellaeaWrightiana
of recent writers.The name mucronata,
asapplied
toany
species of thegenus
Pellaea, datesfrom
1856,with the
publication of Allosorusmu-
cronatus
D.
C.Eaton, based wholly on small specimens
collected 'inthe
hillsnear the bay
ofSan
Francisco, California,"by Major A. B. Eaton. This
material, recentlyexamined,
clearly bears outEaton's
laterstatement!
that his"original
Allosorusmucronatus was founded on small specimens
ofP.
ornithopus, ofHooker" (1858). In the Botany
ofthe Mexican Boundary Survey (1859), however, Eaton had
transferredA. mucronatus
toPellaea,making
Pellaeamucronata include
(inaddition
toP.
ornithopus
Hook.) the two southwestern forms
described in1858
as PellaeaWrightiana Hook, and P. longimucronata Hook.
Recognizing subsequently
the error of associating the distinc- tively Californian plant with the new southwestern
plants,and
objecting (loc. cit.)to
Baker's
use ofthe name mucronata
forthe
latter,Eaton
yet failed totake up the name mucronata
forthe
Californian plant,and on the contrary wrote
Allosorusmucro-
*Publishedwith the permissionofthe SecretaryoftheSmithsonianInstitution.
+ Ind.Fil. 481. 1906.
IFerns N.Amer.2: 8.1880.
44—Pkoc. Biol.Soc.Wash.,Vol.30. 1917. (179)
natus as
a
straightsynonym
of Pellaea ornithopusHook.*
The
lattername, though
singularly appropriate,must
evidentlygive
way
to Pellaeamucronata,
limited to theabundant
Califor-nia plant.
Not much
attentionappears
tohave been given
tothe PellaeaWrightiana and
P.longimucronata
ofHooker, founded
respec- tivelyon
nos.2130 and 2131
ofCharles Wright's New Mexican
collections of1851-52. They have been regarded
asforms
ofa
single species, P. Wrightiana.Actually they
representtwo
species,
and
the charactersby which they may be
distinguished are thosegiven
originallyby Hooker and
restatedby Davenport.f the
latter recognizinglongimucronata only
varietally,however.
Davenport's
varietycompacta appears
tobe
a valid species, allied to P.mucronata
(P. ornithopus).The synynomy and
range,and some
of the distinctive char- acters ofthe
four species, aregiven
below.The rhizome
scales exhibitsuch pronounced
variation innearly
all respects as tobe
of far lessimportance than
isusual,and
charactersdrawn from them
arepurposely omitted.
Pellaea
mucronata
(D. C. Eaton) D. C. Eaton inTorr. U. S.& Mex.
Bound.
Bot. 233. 1859.Allosorusmucronatus D. C. Eaton, Ainer.Journ.Sci. II.
22:
138. 1856.Pellaea ornithopus
Hook.
Sp. Fil. 2: 143.pi. 116. A. 1858.Allosorus ornithopus Kuntze, Rev. Gen. PI. 2:806. 1891.
Type.
—
Collectedon
"hills nearthebayofSan Francisco, California,"by Major A. B. Eaton.
The
labelaccompanying
thespecimen givesthe additional locality ''Monte Diablo."Distribution.
—
Widelydistributed in California,occurring fromMen-
docinoCounty
southward intheCoastRangestoSan
DiegoCounty and Lower
California,and
from Shastaand
Lassen Counties southward through thefoothillsof the Sierra Nevada, ascending to 1,650metersin ElDorado
County, or possibly higher,tAn
extremely variable species, showing the utmost diversity in size,number, and
spacingof segments, accordingtoseason,habitat,andgeo- graphical position.What may
be regarded as thenormal
form is that figured by D. C. EatoninFernsofNorth America,§ butthere aremany
departuresfromthis, the
more
noteworthy being(1)extremelyluxuriant specimens from Los Angeles County, with pinnae alternate,many
of•Op.cit,2: 12.
+Cat.Davenp.Herb. Suppl.45, 46. 1883.
tReportedby Watkins(FernBull.10: 70.1902) toascendto 9.000feetintheSierra Nevada.
§2:pi. i7, Jigs. 7-10.
Maxon —Notes on
Western Species ofPellaea. 181 them more
than 10 cm. long; (2) specimens from thevicinity of Teha-
chapi, in Kern
County, with exceedingly small, numerous, close-set
segments; (3) long-stalked fronds from Lassen County
(Austin 1367)
which suggest P. brachypterabut are readily distinguished from that by
theirvery
much
longersecondaryrachisesand
shorter, smaller pinnules.The
northern specimensshow
atendency toward simple pinnules (sec- ondary pinnae); whereasinspecimens from Whitewater (G. R. Vasey) the largersecondary pinnaeconsist of asmany
as9to11 sessilesegments.Entirelycharacteristicspecimensareat
hand
from northernLower
Cali- fornia {Mearns 3193, 3791); also from SantaCatalina Island,where
the plant is called "tea-fern," from its use for tea. According to A. A.Eaton,
who
hasdiscussedthis speciesveryinterestingly,*it iscalled also"black fern
" and
"poison-fern," thelattername
indicatingitsreputed poisonouseffectupon
sheep.The
frondsof P.mucronataarealmostinvariablyfertile.The
segments are thick, sharply mucronulate,revolute to the middle at all stages of growth,and
often transversely corrugate at maturity, with themargins closely repandand
finelysinuate-dentate.The more
subdivided laminaand
theabsenceofsterilefrondswill atoncedistinguish this speciesfrom the occasional fronds of P. longimucronata with small segments.The
relationshipwith P. compactaisdecidedlycloser.Pellaea
Wrightiana Hook.
Sp. Fil. 2: 142. pi. 115. B. 1858.Pellaea mucronata D. C.
Eaton
in Torr.U.S.& Mex. Bound.
Bot. 233.1859, in part; notAllosorusmucronatus D. C. Eaton, 1856.
Type.—
CollectedinNew
Mexico,in 1851,by
Charles Wright(no. 2130), theexactlocalitynotstated. Accordingto dataintheGray Herbariumt
itwas collected on
"mountains
around the Cobre.On
rocky ledges,October 20."
"Cobre"
is one of thenames
for SantaRita, amining camp
inGrant
County,altitude 1900 meters, about 15mileseast of Silver City.+A
specimenof Wright2130,agreeingcloselywith Hooker'sfigure, is intheU. S. National Herbarium.Distribution.
—
SouthwesternOklahoma and
central Texas to Arizonaand Lower
California.Fronds mostlyfertile, at least in the upper half or two-thirds, occa- sionally wholly sterile; lamina linear or in largewell developed speci-
mens
sometimes lanceolate or narrowly triangular, the basal pinnae reduced or not; pinnaeshort, insome
plantsall buttheupper one trifol.iolate (these gradually simpler),in others themiddle
and
sometimesthe basalpinnae with2or occasionally 3 pairs of pinnules besides the large terminal one, theupper pinnae trifoliolate, finallysimple; pinnules dis- tant, nearly equal, relatively large, semiadnateorsessile,the fertileones closely revolute one-halftoone-third the distanceto the middle, flatfish, themargin entire or nearlyso, slightly altered, evident at theapexasa•FernBull. 12: 113, 114. 1904.
tSee Standley, Contr.U.S.Nat. Herb.13: 175. 1910.
JOp.cit. 170.
narrow
cartilaginousbordertothe short raucro; sterilepinnules similar but broader, plane, the margin pale, thinnish; leaf tissue coriaceous, lightgreen,slightlyglaucous.ReadilydistinguishedfromP. longimucronata Hook., with which it is contrasted below. It has sometimes been distributed
mixed
with P.longimucronata
and
apparentlygrows incompany
with that in severallocalities, butthe rangesthoughsimilar do not coincide, P. Wrightiana extending farther to the east.
The Oklahoma
record is of specimens collected atQuanah Mountain
in1891 byC. S. Sheldon(no.192), listed*asP. ternifolia.
The
central Texas specimens are from Llanoand
Gil- lespieCounties (Bray303, Reverchon 1216,Jermy
351),and
the other Texas plants arefrom theChenateand Cornudas Mountains
(Havard).In
New Mexico
ithas beencollected in the Burro Mountains byRusby and
intheOrgan
Mountains byWooton and
Standley,and
has been re- collected at the type localityby
J.M.
Holzinger, August 27-September12, 1911. There are
numerous
Arizona specimens,coming
from several parts of the State but largelyfrom themountain
rangesofPima and
Cochise Counties,and
thereisasingle specimen fromLower
California (ElRancho
Viejo, April29, 1889, T. S.Brandegee).The
speciesascends to at least2,400metersin theChiricahua range (Blumer 1867).Pellaea
longimucronata Hook.
Sp. Fil. 2: 143. pi. 115. A. 1858.Pellaea Wrightiana longimucronata Davenp. Cat. Davenp. Herb. Suppl.
46. 1883.
Pellaea truncataGoodding, Muhlenbergia
8
:94. 1912.Type.—
CollectedinNew
Mexico,in 1851, byCharlesWright
(no.2131), theprecise localitynotgiven. According to a note by Wooton, accom- panying a specimen of thisnumber
in theNational Herbarium, itwas
collectedon
"mountain
sidesnear Conde'sCamp,
Sept.1, 1851,"Conde'sCamp
being in the western part ofGrant
County, near the Arizona boundary.The
specimen accords with Hooker's descriptionand
illus- tration.Distribution.
—
SouthwesternNew Mexico
to southern Utah, extreme southern Nevada,and
western Arizona;also insouth-central Colorado.Fronds mostlyfertile, but sterile fronds
much commoner
than in P.Wrightiana, partiallyfertileonesless
common
; laminatriangular-ovate, fully bipinnatealmostto theacuminate simplypinnateapex; middleand basal pinnae of fertile fronds the largest, the pinnules (6 to10pairs) rigidly divaricate, small, usually decreasing in size toward the reduced apical segment, all sessile or short-petiolate, articulate,many
ofthem
deciduous after maturity; fertile pinnules sharply mucronate, very strongly revolute (the edges frequently meeting), the marginsslightly modified, paler, usuallysomewhat
erose-denticulate; sterile pinnules fewer, larger, relatively broader, often truncate, conspicuously long- mucronate, themargins plane or repand, erose-denticulate or sometimes*Contr.U.S.Nat.Herb.1:201.1892.
Maxon —Notes on
Western Species ofPellaea. 183
irregularly short-spinulose; leaf tissue extremely coriaceous, grayish
green, pruinose.
Pellaealongimucronata differs from P. Wrightiana in having the lamina broader
and
trulybipinnate nearlythroughout,trifoliolatepinnae mostly wanting; in itssmallerand more numerous
pinnules, thesemore
rigid,
more
slenderly mucronate,and
distinctly articulate, beingmore
or jess deciduous with age from the short knob-like petiolules;and
in its harsher,much more
coriaceous texture, the pinnulesmore
strongly revo- luteand
having an altogethercharacteristicgrayish-pruinoseappearance.These characters, though previously noted, have not recently been re- garded as of importance. Yet they were regarded as sufficient forthe establishmentof
two new
speciesby
soconservative a botanistasHooker,who
had, moreover, only very scant material;and
in the greatnumber
of specimens
now
athand
they areso easilymade
outand
sounmistaka- ble as to leaveno doubtofthespecificdistinctnessofthetwo
forms.The
supposed intermediatestatesmentioned by Eaton have not beenseenand
certainly are notfound in hisown
herbarium,hisspecimens beingdefi- nitelyreferabletoonespeciesorthe otherinaboutequalnumber.
Pellaea longimucronata is apparently very
much commoner
than P.Wrightiana. It is abundantly represented
from New Mexico
in the collections ofWooton and
Standley in theOrgan
Mountains,and
ofRusby
intheBurro Mountains. Itiswidely distributedinArizona,being especiallycommon
apparentlyinthe Santa Catalina Mountains, here as intheOrgan
Mountainsand
elsewhere occurring in association with P.Wrightiana; the typeofP. truncata isfromthe
Mule
Mountains.The
Colorado record is of specimens collected atCanyon
City,Fremont
County, by Brandegee,*and
thisis the onlylocalitygiven inRyd
berg's FloraofColorado; itissubstantiated byaspecimenintheEaton
Herba- rium.The
singleUtah
specimen athand
is from SilverReef, altitude about 1,050 meters, Jones 5,149aq.The Nevada
specimensaretwo: Vir- gin River, Bunkerville,Goodding737;Mica
Spring,altitude1,200meters, Jones5,055.The
lastcited specimen includes afrond showingthe proxi-mal
basal pinnule of the basal pinnaefully pinnate,withthree distinct segments.Pellaea
compacta
(Davenp.)Maxon,
sp. nov.Pellaea Wrightiana compacta Davenp. Cat. Davenp. Herb. Suppl. 46.
1883.
Rhizome
woody, multicipital, the branches nodose, short-creeping, 1 to 2cm. long, about 1 cm. thick, freelyradicose beneath, densely palea- ceous, the scalesdarkbrown
in mass, forming a close tufted covering, linear, long-attenuate, falcate or straight, the characteristic larger ones 5 to 7mm.
long, 0.5 to 0.7mm.
broad, with a broad, thick, strongly sclerotic, dark brown, lustrousmedian
band, the borders thin, pale, transparent, plane or distinctly repand, conspicuouslyand
irregularly denticulate; fronds numerous,closely fasciculate, mostlyfertile, 20to35•Port.
&
Coult.Syn.Fl.Colo.153. 1874.cm. long, the thick glossydark
brown
flexuousstipesmostlymuch
longer thanthe lamina, glabrous; lamina narrowlyoblong, acuminate, 9 to 13 cm.long, 2.5 to 4.5cm. broad, bipinnate; pinnae about 15 pairs,usually oblique in drying, the basal pair distant, those above gradually closer, thoseoftheapical portion contiguous or imbricate, gradually or abruptly simple; pinnules of the larger pinnae 5 to 7 pairs, simple, nearly uni- form,mostlyclose,sessileor very short-stalked,with a short cartilaginous mucro, thesterileones oval or oblong, plane, punctate beneathand with themargins minutelyerose; fertilepinnules narrower, broadlyrevolute, often conduplicateand
falcate with age, transversely wrinkled, the slightly thinner margins repandand
finely sinuate-dentate; sporangia borne in a broad marginal zone, almost wholly concealed; leaf tissue rigidly spongiose-herbaceous, the surfaceslightor grayish green.Described originally
by
Davenport as "a remarkable distinct form with longstipites
and
denselycrowded compactpinnules, collected intheSan
Bernardino Mts. ofCaliforniabyW.
G. Wright." Thiscollectionis notrepresented in the National Herbarium, but there is a specimen in theEaton Herbarium, collected in 1879and
ticketedin Wright'shand
as"
gathered
on
Mt.San
Bernardinoatan
altitude of 7,000 ft.," which isundoubtedlytypical. Itis
marked
by Eaton"
var.californica
Lemmon,"
a
name
apparently unpublished,and
agreeswithanotherEaton specimen, underthesame
name,marked
"San
Jacinto Mt., Parish brothers, 1880."AccordingtoParish* the plantis "frequent in the San Bernardino
and San
Jacinto Mts.,growinginstonysoilon
dryslopes at6,000to 8,000ft.altitude,"
and "has
beencollectedby
Brandegee in theProvidenceMts., intheMojave
Desert."The
followingspecimensareintheNational Herbarium:Among
rocksonsteepsides ofSanJacintoMountain, July, 1881, S. B.
& W.
F.Parish 511;Snow
Canyon,San
Bernardino Co.,alt. 1,800 meters,June
20, 1901,S. B. Parish 5050; stonyridges,
San
Antonio Mountains, July 11,1902,Abrams
2684; south slope, Sugarloaf, San Bernardino Mts.,alt. 2,400 meters,forming"denselow clumpsin slides,"July 11,1906, J.& H. W.
Grinnell215.
Pellaea compacta is allied to P. mucronata, agreeing in
most
general characters, butdiffering sufficiently initssimple, lesssharplymucronate pinnules,itscongestedhabit,itslong-stipitate fronds,and
itsmore
broadly striped rhizome scales.With
P. Wrightiana ithasnocloseconnection whatever,and
fromP. longimucronataitdiffers widely in its lessersize, itscongested pinnae,and
its close, very short-pointed fertile pinnules, these not only revolute to the middle butcommonly
conduplicateand
distally falcate. Apparently P. compacta grows in
company
with P.mucronata,
and
itwould
be interestingto studytheir relationshipinthe field.*FernBull.12: 8.1904.