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 AECIAL STAGE OF PUCCINIA OXALIDIS
W. H. LONG AND R. M. HARSCH

In JulyI 9 I 5 the juniorwritercollectedan undescribedA ecidium

on the leaves of Berberisrepens in Bear Canyon, located in the
Sandia Mountains about I 5 mileseast ofAlbuquerque,New Mexico.
At the time the Aecidiumwas discoveredno clue was found as to
what hosts might harbor the alternate stages of this rust. The
marked differencesin the microscopic characters of the new
Aecidium easily separated it fromthe aecial stage ofboth Puccinia
graminisand P. koeleriae,the only two otherrustsknownto occur
on species of Berberis. The firstassumption was that this new
rustmighthave its alternatestage on some graminaceoushost,but
carefulfieldwork in the springof i9i6 by the senior author soon
dispelled this theory,since this Aecidium was often found abun-
dantlyin localitieswheretherewere no possiblegrass hosts. Field
observationsshowed that this rust always occurred in localities
where plants of Oxalis violacea and Berberisrepens were closely
associated,and whentheywerenot associated no rustwas foundon
the Berberis. Later in the springof i9i6 the seniorwriterfound
young leaves of Oxalis violacea bearing the primaryurediniospores
of Puccinia oxalidisin directcontact with the old aecia whichhad
sporulated. This association of the two rusts was constant
throughoutthe canyons in the Sandia Mountains, where the two
hosts occurredin proximityto each other,while neitherrust was
found on eitherhost when the hosts werewidelyseparated.
With this positivefieldclue as a guide, inoculationswere made
at Tejano ExperimentStation, in the Sandia Mountains, about
30 miles fromAlbuquerque, on io wild plants of Berberisrepens
growingin the open. Afterthoroughlywettingthe plants, living
old leaves of Oxalis bearing germinatingteliosporesof Puccinia
oxalidis were placed above young leaves of Berberis. Both inocu-
lated and checkplants wereprotectedby placing tincans overthem.
These inoculationswere made September20, I9I6. The tin cans
were removed September 23. On October 20 the io inoculated
475] [Botanical Gazette, vol. 65
476 BOTANICAL GAZETTE [AIAY

plants were examined and all of them were found to be infected;

some so badly that the leaves were dying,while all (6) of the check
plants were healthy. At this date pycnia only were present,
exuding droplets of a sweetish sticky fluid like honey-dew.
Another trip was made to the Station June 30, I9I7, when the
inoculated leaves showed well developed aecia, while the check
plants were still free of the rust. These inoculations were not
considered absolutely conclusive, however, since the Berberis
plants inoculatedwerein the open and thereforesubject to external
contamination.
In the fallof i9i6 bulbs of Oxalis violaceawere transferred
from
the mountainsto Albuquerque, a distance of some I5 miles from
any Berberisplants, and thereforefree from any possible exter-
nal contamination. Fresh but non-sporulatingmaterial of the
Aecidium on Berberiswas obtained from Bear Canyon June 22,
I9I7. At this time no Oxalis plants had appeared above ground
in the vicinityof the infectedBerberisleaves in the Canyon, but
the Oxalis plants transferredto Albuquerque the preceding fall
werein fullleaf. The infectedBerberisleaves were moistenedand
kept overnightunder a bell jar to start sporulation. On June 23
two species of Oxalis (0. violacea and 0. stricta)were inoculated
undercontrolconditionswith the aeciosporesfromBerberis. Bell
jars were kept over the plants 6o hours. Checks were also made.
On June 30 many of the inoculated leaves of 0. violacea had the
typicalurediniaof Puccinia oxalidis,while the check plants as well
as all plants of 0. strictawere freeof the rust. July 22 telia were
presenton the inoculated leaves of 0. violacea. The inoculations
herereported,togetherwith those made at the Tejano Experiment
Station, prove conclusivelythat the new Aecidium on Berberis
repensis the alternatestage of Puccinia oxalidis, a descriptionof
whichis herewithgiven.

PUCCINIAOXALIDIS (Lev.) Diet. and Peck

0. Pycnia amphigenous but mainly epiphyllous, seated on
pallid to slightlyreddishspots 4-8 mm. in diameter,conspicuous,
conic-globoid,honey-yellowbecoming blackish brown, appearing
in the fall of the year when the pycnosporesare dischargedin a
i9i8] LONG o HARSCH-PUCCINIA OXALIDIS 477

sweetish sticky liquid. In the spring when the aecia appear the
pycnia are blackish brown.
I. Aecia hypophyllous,seated on pallid to reddishbrownspots
which later become dark brown, crowded in irregularannular
groups 4-8 mm. across, aecia orange color when fresh,cylindrical,
I-I. 25 mm.highbyo. I5to 0.2 mm. indiameter, peridium opening
at apex very irregularly,very slow to open and very tough, seg-
ments slightly if at all reflexed,usually falling away piecemeal,
peridial cells not overlapping,in face view irregularlyoblong to
polygonal, IO-I 7 XI7-27 g, in side view pulvinate I4-I 7 X 20-24 Y,
innerwall verruculose2-2 .5 g thick,outer wall irregularlystriate,
3-4 A thick,walls colorless, content of cells orange; aeciospores
irregularlyoval, ovate to subglobose, angular, IO-I3 XI3-I7 A,
average forten, II XI4.4 A; walls colorless,faintlyverruculoseto
smooth,I .5-2 g thick,pores indistinct.
On Berberidaceae: BerberisrepensfromNew Mexico as follows: Bear
Canyonby R. M. Harsch,July7, 1915 (no. 5554);' by Bartholomew and Long,
June22, I917 (no. 628i), materialused forinoculatingOxalis violaceaplants;
by Long,August2, I9I7 (no. 6284); Tejano ExperimentStation,by Longand
Seay, Juneand Julyi9i6 (nos. 6005, 6oo6, 602I, 6097); by Long,JulyI917
(no. 6285), materialobtained by inoculatingBerberisplants September20,
i9i6, with teliosporesof Puccinia oxalidis fromOxalis violacea July I9I7
(no. 6286).

II. Uredinia hypophyllous, subepidermal, in irregular to

orbicular groups 2-6 mm. across, often confluentand covering
entire surface of leaf, round, O. I-0.3 mm. across, soon naked,
at firstorange buffand waxy, later fadingsomewhatand becom-
ing pulverulent,rupturedepidermisinconspicuous; urediniospores
globoid or elliptical globoid, I5-20 X I7-25 g; walls thin, about
I g, minutelyechinulate,germpores uncertain.
III. Telia hypophyllous, in orbicular to irregular groups
2-5 mm. across, often confluentover entire leaf surface,subepi-
dermal,rupturedepidermisinconspicuous,soon naked, orangebuff,
waxy, round, 0.1-0.3 mm. across. Teliospore ellipsoid to oval,
12-22XI7-28 gA,rounded or obtuse at both ends, slightlyor not at
all constrictedat septum; septum often oblique; walls colorless,
IAll herbariumnumberscited in this article referto the herbariumnumbersof
theseniorwriter.
478 BOTANICAL GAZETTE [MIAY

smooth, thin, less than i g thick; pedicel colorless,thick, about

as long as spore.
On Oxalidaceae: Oxalis violacea fromNew Mexico as follows: Albu-
querque,by Long,July1917 (no. 6282), materialobtainedby inoculatingwith
aeciosporesfromBerberisrepensJune 20, I9I7; Tejano ExperimentStation,
by Long and Seay, Julyand Septemberi9i6 (nos. 6014, 6ioo, 6102). Also
reportedon followinghosts: fromJamaica,Oxalis martiana; fromMexico,
0. vallicola,Oxalis sp.;
0. trinervis,
Oxalis divaricata,0. latifolia,0. tetraneuris,
fromTexas, 0. violacea; fromBrazil,0. neuwiedii.
The roestelia-likeaecia and othercharactersof thisrustindicate
its relationshipin a general way to the genus Gymnosporangium,
while some of its characters show affinityfor the genus Eriospo-
rangium,fromwhich,however,its very tough,persistentperidium
would excludeit. It does not belong to the genusA rgomyces, where
ARTHUR has provisionally placed it. If one were following
ARTHUR'S nomenclature,the rust would probably belong to a new
genus,but the writerspreferto leave it underthe old genusPuccinlia
forthe present.
OFFICE OF INVESTIGATIONS IN
FOREST PATHOLOGY, BUREAU OF PLANT INDUSTRY
ALBUQUERQUE, N.M.