Amor. Jour. Bot. ;";1(1): 1-(;.19(;4.

CHROMOSOME NUMBERS IN NORTH AMERICAN LORANTHACEAE:

(ARCEUTHOBIUM, PHORADENDRON, PSITTACANTHUS,
STRUTHANTHUS) l.~
DELBERT WIENS
Department of Biology and Institute of Arctic and Alpine Research, Universitv of Colorado,
Boulder, Colorado

ABSTHACT
Chromosome numbers are reported from (;7 populations of 3(; taxa, mostly in Phoradendron,
The bask number is 14 in Phoradcndron and probably also in ..Irreuthobiuni. The 4 species of
Struiluuuh.u« for which chromosome numbers are recorded suggest that the basic number is 8. Num-
bers of n = 8 and n = 10 have been reported for Psitlacanth.un, In Phoradcndron. the 22 taxa
examined are all diploid, although 1 instance of polyploidy was discovered. Objects interpreted as
supernumerary chromosomes were discovered in a number of species of Phoradcrulron; no evidence
of sex chromosomes previously reported in Phoradendron. was observed. The Lorunthaooae have
chromosomes comparnhle in size to the largest in the plant kingdom.

I'm; LOnAN'l'HACEAE is composed essentially of
woody, pan-tropical plants parasitic on the aerial
portions of vascular plants. Some genera, however,
have representatives extending well into the
temperate regions, e.g., Phorodendrcn and Vise1l1n.
Arccuihobium. occurs essentially in the temperate
regions of the northern hemisphere. Psitiacanihus
and Struihanthus are restricted primarily to the
American tropics.
The known information on chromosome num-
bers in the Loranthaceae has been summarized
by Gill and Hawksworth (19Gl), who list the
chromosome numbers of 22 species occurring in
11 genera. The family contains approximately
1,100 species and perhaps :30 genera (Lawrence,
HJ;"j1). In the present study ()7 chromosome
numbers are reported for ;H species in 4 genera.
Heretofore, Phoradendron serotinum (= Ph. flaoe-
secns) appears to be the only member of the
family whose chromosomes have been studied in
detail (Baldwin and Speese, 1%7).
In Phoraderulron; microsporogenesis begins ap-
proximately 2-:j months before flowering and is
generally completed by anthesis; however, in rare
instances dividing pollen mother cells may still be
found in the youngest buds of the inflorescence
when the first flowers open. In the species of
Arccuihobium. studied, the pollen mother cells
begin to divide approximately 1 month before
anthesis, except in A. americanum, which under-
goes meiosis during the summer and flowers in
the succeeding spring. Mierosporogenesis probably
continues over a period of about :1 weeks. The
1 Received for publication Januarv 1 J, 1!Hi3. Based on a
portion of a dissertation submitted to the Claremont
Graduate School in partial fulfillment of the requirements
for the degree of Doctor of Philosophy.
2 This work was supported, in part, by a grant from the
society of Sigma Xi.
[The .TOURNAL for November-December (50: 965-10(1) was issued November "27, 1!J63]
AMERICAN .JOURNAL OF BOTANY, Vol, 51: No.1, 19(;4
approximate dates at which meiosis occurs in the
species examined may be determined from Table 1.
One anomalous situation was discovered in the
typical form of A. uaqinaium occurring in central
Mexico, Staminate material of this form was
undergoing meiosis in August; however, in Color-
ado forma cnjpiopodum undergoes meiosis in
March,
Mm-nons-e-Chromosomc numbers were deter-
mined in microsporoeytes;" except in 2 instances
where dividing tapetal cells were utilized. Material
was fixed in ;1: 1 absolute alcohol and acetic acid.
Anthers were squashed and stained in acetocar-
mine. The chromosomes stain deeply and rapidly
and there is no cytoplasmic staining.
A list of species for which chromosome numbers
were determined and the collection data are
summarized in Table 1. The nomenclature em-
ployed in Phoraderuiron. and Arceuthobium is that
of Wiens (1DG4) and Gill (Hl;jij), respectively.
Voucher specimens are deposited in the Rancho
Santa Ana Botanic Carden Herbarium. The
drawings of chromosomes were prepared with the
aid of a camera lucida.
lb,suvfs AND DISCUSSION-The chromosome
numbers determined in Phoradendron are, with 1
exception, n = 14. The chromosome number of
the species of A rcciuhobium examined also was
determined to be n = 14, and no species with
n = l;j, as reported by Pisek (1D24), were found.
The numbers observed in A rceuihobium. are in
agreement with those of Dowding (1D;n) who
first reported n = 14 in A. americanurn. The
chromosome numbers determined in Struihanthu«
(n = 8, 2n = Hi) are the same as those reported
by Covas (lD49) for the South American species,
S. angust~folius (Griseb.) Haum,
3 I am indebted to Drs. William M. Klein and T. Paul
Maslin for making several collections of buds.

1
2 AMERICAN JOURNAL OF BOTANY [Vol. 51

TABLE 1. Chromosome numbers in Arceuthobium, Phoradendron, Psittacanthus and Struthanthus.

Species Chromo no. Supern. chromo Voucher collection data

A. americanum Nutt. ex Engelm. n = 14 COLORADO. Boulder Co.: near Nederland.

Wiens 2966. 8/7/G1.
A. campylopodum Engelm. forma campylo- n = 14 CALIFORNIA. San Bernardino Co.: Big
podum Bear Lake. Wiens 2741. 7/9/GO.
A. campylopodum f. cyanocarpum (A. Nelson) n = 14 COLORADO. Boulder Co.: Ward. Wiens
Gill 2926. 7/6/61.
A. campylopodum f. divaricatum (Engelm.) n = 14 CALIFORNIA. Inyo Co.: Whippoorwill
Gill Flats, on Big Pine-Saline Valley Rd.
Wiens 27.40- G/18/60.
A. vaginatum (Willd.) Presl. f. vaginatum n = 14 MEXICO. Veracruz: south of Perote. Wiens
(Fig.2B) 2574.8/18/59.
A. oaqinaium f. cryptopodum (Engelm.) Gill n = 14 COLORADO. Boulder Co.: near Boulder.
Wiens 2802. 3/23/G1.
Ph. bolleanuni (Seem.) Eichler, subsp. n = 27 CALIFORNIA. Sonoma Co.: ncar Monte
densum (Torr.) Wiens Rio. Wiens 2723. 6/14/GO.
Ph. bolleanum subsp. densum n= 14 Five populations"
Ph. bolleonum subsp. pauciflorum (Torr.) n = 14 2 CALIFORNIA. Los Angeles Co.: Mt. Baldy
Wiens ski area, San Gabriel Mts. Wiens 2461.
5/27/59.
Ph, brachystachyum (DC.) Nutt. n 14 MEXICO. Morelos: lava fields north of
Coajomulco on toll rd. to Mexico City.
Wiens 2530. 8/15/59.
Ph. colijornicum Nutt. n = 14 2-8 CALIFORNIA. San Bernardino Co.: near
Barstow. Wiens 2662. 10/14/59.
Ph. calyculatum Trclease n= 14 MEXICO. Chiapas: IG miles se. of San
Cristobal. Wiens 2976. (coli. by T. Paul
Maslin, 7/23/Gl).
Ph. capitellaiuni Torr. ex Trelease n 14 ARIZONA. Yavapai Co.: near Sedona. Wiens
2763. 9/13/60.
Ph. qoleotiii Trelease n 14 MEXICO. Puebla: ncar Zacatecas, w. of
Perote on hwy. 140. Wiens 2581. 8/18/59.
Ph. quazumae Trelease n = 14 MEXICO. Sinaloa: hwy. 15, n. side of
Mazatlan. Wiens 2495. 8/11/59.
Ph. [uniperinuni Engelm. subsp. juniperinum n = 14 Nine populations':
Ph. juniperinum subsp. libocedri (Engelm.) n = 14 CALIFORNIA. El Dorado Co.: Institute of
Wiens Forest Genetics, Placerville. Wiens 2732.
G/17/GO.
Ph. [uniperinum subsp. libocedri n 14 CALIFORNIA. El Dorado Co.: near Kyburz,
hwy. 50. Wiens 2734. 6/17/GO.
Ph. [uniperinum subsp. libocedri n 14 CALIFORNIA. San Bernardino Co.: Stock-
ton Flat, Lytle Creek Canyon. Wiens 2484.
G/17/fi9.
Ph. lanaium Trelease n 14 3-4 MEXICO. Queretaro: Ileal' Cadereyta. Wiens
2595. 8/20/59.
Ph. minulijolium Urban n = 14 MEXICO. Veracruz: ca. 3 miles S. of Perote.
Wiens 2578. 8/18/59.
Ph. nervosum Oliver n 14 MEXICO. Chiapas: 2G miles so. of San
Crist6bal. Wiens 2080. (colI. by T. Paul
Maslin, 7/2G/61).
Ph. puberuluni Trelease n 14 2 MEXICO. Nayarit: ca. 20 miles s. of Topic
on hwy. 15, Wiens 2506.8/12/59.
Ph. puheruluni n 14 MEXICO. Jalisco: ncar Magdalena on hwy,
15. Wiens 2511.8/13/59.
Ph. reichenbachianum Oliver n 14 MEXICO. Morclos: ncar Temple of Tcpox-
tlan, Tepoxtlan. Wiens 2531. 8/15/59.
Ph. robinsonii Urban n 14 2-3 MEXICO. Puobla: near Tehuac.in. Wiens
2551. 8/17 /.~9.
Ph. schumannii Trelcasc n 14 MEXICO. Durango: hwy. 40, ncar km. 10LO
btwn. Durango and El Saito. Wiens 2632.
8/23/59.
Ph. tomenlosum (DC.) Engelm. subsp. n 14 Four populationss
tomeniosum.
Ph. tomeniosum subsp. macrophsflluni n 14 Five populations"
(Engclm.) Wiens
January, 1964] WIENS--LORANTHACEAE

TABLE I.-Continued

Species Chromo no. Supern. chromo Voucher collection data

Ph. uelutinuni (DC.) Nutt. n = 14 3 MEXICO. Mexico: ca. 10 miles e. of Toluca

on hwy. 15. Wiens 2529. 8/14/59.
Ph. uelutinuni n = 14 MEXICO. Puebla: ca. 20 miles n. of Tehuacrin
on hwy. 150. Wiens 2553. 8/17/59.
Ph. uillosum (Nutt.) Nutt. subsp. uillosum. n = 14 Eight populations"
Ph. oillosuni subsp. coryae (Trelease) Wiens n = 14 ARIZONA. Coconino Co.: Oak Creek Can-
yon. Klein 1470. 7/26/60.
Ph. oillosum subsp. coryae n = 14 NEW MEXICO. Socorro Co.: rd. to South
Baldy Peak, Magdalena Mts. Wiens 2661.
8/21'/59.
Ph. villosum subsp, flacum (1. M. Johnston) n = 14 MEXICO. Durango: km. 995 btwn. Durango
Wiens and EI Saito on hwy. 40. Wiens 2625.
1'/23/59.
Ph. »illosum subsp. flauum n = 14 MEXICO. Durango: km. 1010 btwn. Dur-
ango and EI Saito, on hwy. 40. Wiens 2630.
8/23/59.
Ps. americanus (Jacq.) Mart. (Fig. 2A). n = 8 MEXICO. Mexico: Tepetlixpa. Wiens 2538.
8/16/59.
Ps, calsjculaius (DC.) D. Don 2n = 16 MEXICO. Sinaloa: hwy. 15, ca. 30 miles s, of
Navajoa. Wiens 2491. 8/10/59.
S. inconspicuous (Benth.) StandI. n = 8 MEXICO. Sinaloa: hwy. 15, ca. 40 miles n.
of Mazathin. Wiens 2494. 8/ll/59.
S. micropliullus (HBK) D. Don (Fig.2C). ti = 8 MEXICO. Michoacan: hwy. 15, near Santa
Rosa, ca. 50 miles e. of Morelia. Wiens
2522.8/14/59.
S. »enetus (HBK) Blume 2n = 16 MEXICO. Sinaloa: hwy. 15, near San
Lorenzo, s, of Culiacrin, Wiens 2498.
8/11 /59.

• CALIFORNIA. Inyo Co.: Whippoorwill Flats, on Big Pine-Saline Valley Rd. Wiens 2739.6/18/60. San Bernardino
Co.: near junct. of hwy. 138 and Big Pines Rd. Wiens 2681.4/30/60. (Fill:. 2E). ARIZO~A. Yavapai Co.: near Sedona.
Wiens 26.91.5/6/60. Gila Co.: near Pine. Wiens 2706.5/7/60. MEXICO. Coahuila: ca. 40 miles s. of Saltillo on hwy. 57.
Wiens 2607. 1'/21/59.
b CALIFORNIA. El Dorado Co.: hwy. 89 ca. 5 miles s, of junct. with hwy. !i0. Wiens 2785.6/17/60. Alpine Co.: near
Monitor Pass. Wiens 2786. 6/17/60. Inyo Co.: Whippoorwill Flats, on Big Pine-Saline Valley Rd. Wiens 2788. 6/18/60.
San Bernardino Co.: Big Bear Lake. Wiens 2485.6/25/59. ARIZONA. Coconino Co.: Ashfork. Wiens 2839. 5/25/61.
Coconino Co.: Sedona. Wiens 2697 ..5/7/61. Yavapai Co.: 13 miles s. of Camp Verde on rd. to Pine. Wiens 2714. 5/7/60.
Pin'a Co.: rd. to Mt. Lemmon. Wiens 2703.5/7/60. NEW MEXICO. Taos Co.: ncar Taos. Klein 1483.7/27/60.
c MEXICO. Durango: ncar La Cuesta on hwy. 4!i btwn. Durango and Parral, Wiens 2646.8/25/59. Durango: ca. 20
miles w. of Le6n Guzman on hwy. 40. Wiens 2620. 8/22/59. Coahuila: ca. 50 miles e. of Torreon on hwy. 40. Wiens 2616.
8/22/!i9. Coahuila: ca. 28 miles e. of Saltillo on hwy. 60. Wiens 2613.8/21/59.
d CALIFORNIA. Los Angeles Co.: Cobal Canyon, near Claremont. Wiens 2761. 9/3/60. San Bernardino Co.: Cajon
Pass eamporound. Wiens 2668. 10/14/59. ARIZONA. Coconino Co.: Sedona. Wiens 2764.9/13/60. TEXAS. EI Paso
Co.: near Anthony. Wiens 2660.8/28/.59. MEXICO. Coahuila: Le6n Guzman, w. of Torre6n on hwy. 40. Wiens 2617.
8/22/59.
p CALIFOR!'\IA. Butte Co.: ncar Paradise. Wiens 2726.6/15/60. Marin Co.: ca. 3 miles n, of San Rafael on hwy. 101.
H1'en' 2722.6/14/60. EI Dorado Co.: ea. 15 miles w. of Placerville on hwy. 50. Wiens 2781.6/17/60. EI Dorado Co.:
r-oar Kyburz on hwy. !i0. Wiens 2738. 6/17/60. San Luis Obispo Co.: ca. 10 miles e. of Paso Robles on hwy. 466. Wiens
2,20. (l/12/CO. Kprn Co.: ncar Frazier Mtn. Park. Wiens 2718. 6/12/60. Los Angeles Co.: Cobal Canyon, near Clare-
mer t. Wiens 2716.5/29/60. San Bernardino Co.: near Running Springs, City Creek Rd. Wiens 2742.7/10/60.

In Psittacanthus, numbers of n = 8 and 2n = 16 Trelease (1916), in his monograph of Phoraden-

were recorded. This differs from the report by
Covas and Schnack (1946) that n = 10. Their
material (Ps. cunefolius Blume I, however, was
also of South American origin. King (1961), in a
study of 2 Mexican species, Ps. auriculatus
(Oliver) Eichler and Ps. calyculatus (DC.) D. Don,
also reported a number of n = 10. The latter
species is the same in which I determined that
2n = If>. More recently, however, King (pel's.
comm.) has also observed numbers of n = 8.
dron, subdivided the genus into 2 primary divisions
without definite rank: (1) the "Aequatoriales"
(with primarily a tropical distribution I, possessing
cataphylls, generally glabrous, and sometimes
monoecious; and (2) the "Boreales," (occurring
predominantly in more northerly temperate
regions), without cataphylls, often pubescent, and
always dioecious, Four species analyzed in this
study, Ph. guazumae, Ph. nertosum, Ph. reichen-
bachianum and Ph. schumanii, are members of
4 AMEHICAN JOURNAL OF BOTANY
the "Aequatoriales," but only Ph. guazumae is
monoecious. All possess the same chromosome
number characteristic of the "Boreales," i.e., 1-1
pairs.
Polyploidy-One polyploid individual (n = 27)
of Ph. bolleanum subsp. densum (Table 1) was
discovered on Cupressus in the Coast Ranges of
northern California. This mistletoe occurs also on
other Cuprcssus stands in the coast ranges and
foothills of the Sierra 1\ evada of California. Addi-
tional work will be necessary to clarify the cyto-
logical condition of these populations, since Ph.
bolleanum subsp, densum is diploid in other parts of
its range (Table 1). Juniperus is the predominant
host of this mistletoe from southeastern Oregon
to northern Chihuahua, but in central Arizona it
again parasitizes Cupressue; however these popu-
lations are also diploid (Table 1).
Supernumerary chromosomes-Structures inter-
preted as supernumerary chromosomes were dis-
covered in Phoradendron (Fig. 1). The number
and systematic distribution of these chromosomes
are listed in Table 1. Their staining properties
appear to be heterochromatic. The number present
in dividing pollen mother cells is variable, but the
mode is 2. Supernumerary chromosomes were not
observed in the other genera examined.
The highest number of supernumeraries noted
(ca. 8) occurs in Ph. calijornicum, This species is
I a cursory review of other plants with large chromo-
FIG. I.-Putative supernumerary' chromosomes at
prometaphase II in Ph. bolleanuni subsp. pauciflornm,
XIOOO.
[Vol. ;')1
restricted to the Sonoran region and is subjected
to greater extremes in aridity than most mistle-
toes. Muntzing (1!l58) and Mooring (J9nO) report
that frequencies of these chromosomes increase in
populations occurring in arid regions and suggest
that they possess some adaptive significance.
More information on the frequency of these
chromosomes in this and some other species would
be desirable. For example, Ph. tomeniosum. subsp.
macrophullum (=Ph.jlal'escensvar. macrophyllum)
is partially sympatric with Ph. calijornicum; but
in other parts of its range, viz. the central valley of
California, it occurs under somewhat more mesic
conditions. The frequency of supernumerary
chromosomes in different parts of the range of
this species could be of interest.
Billings' sex-chromosome hypothesis-Billings
(J 9:32) reported a unique sex-determining mech-
anism for higher plants (an XO type) in Phora-
dendron. His study was based on 2 dioecious
species, Ph. villosum subsp. oitloeum and Ph.
tomentosum subsp. macrophyllum occurring in
southern California. He counted an even number
of chromosomes in the somatic cells of pistillate
plants (2n = 20). However, in the vegetative cells
of staminate individuals he described the presence
of another chromosome which he interpreted as a
sex-chromosome. At metaphase I of microsporo-
genesis he recognized 1 l associations, one repre-
senting a univalent. At anaphase of thehrst
division, 11 chromosomes moved to one pole and
10 to the other. Later Billings (J 9::l:·n examined the
chromosome numbers during megasporogenesis
and reported a consistent number of n = 10.
Baldwin and Speese (1!l!i7) questioned the
validity of Billings' interpretation on the basis of
their cytological work on the eastern mistletoe,
Ph. seroiinum: Current work on the same species
which Billings studied shows that n = 14, and no
structures resembling univalents were detected
during meiosis; however, no pistillate material was
examined cytologically nor have chromosome
numbers been determined in the somatic tissues
of either male or female plants of these species.
No attempt has been made to correlate the
putative supernumerary chromosomes with a
present or past sex-determining function.
Chromosome size-Stebbins (1950) stated that
the genera Trillium and Paris have the largest
chromosomes in the plant kingdom. Textbooks of
cytology, e.g., Sharp (194;·n and Swanson (J !);,,)7) ,
have credited the monocotyledons with having
generally larger chromosomes than dicotyledons.
The dicotyledonous genus Paeonia, however, has
been recognized as an exception. Turner (1!);")8)
reported large chromosomes in Krameria and gave
somes. Covas and Schnack (1!)46) suggested that
Peiuacanthus possessed the largest chromosomes
in the angiosperms. Baldwin and Speese (1\)57)
first demonstrated the large chromosomes in
Phorodendron, The metaphase-I associations of
January, H)(H] WIENS-LORANTHACEAl-J 5

't
'I"~
eA .-.,
11!"
.. ~~
o ." F

~, 4~~
G I 20jJ. I H
Fig. 2. Comparative size of chromosomal associations at metaphase I in plants with large chromosomes.-Fig. 2A.
Psittacanthus alllericanus.-Fig. 2B. Arceuihobium oaqinoiuni forma vaginat1l1l/..-Fig. 2C. Siruihantlius microphsjllusr-«
Fig. 2D. Paeonia calijornica (2 chains of 4 plus 1 bivalent).-Fig. 2E. Phoradendron. bolleanu.ni subsp. densum.-Fig. 2F.
Krameria grayi.-Fig. 2G. Tradescantia paludosa.-Fig. 2H. Liliuui michiganense.-FitJ;. 21. Trillium ooatum, All figures
X ()7ii.
6 AMEmCAN JOURNAL OF BOTANY
some plants recognized as having large chromo-
somes are contrasted with some loranthaceous
genera in Fig. 2. All mounts were prepared accord-
ing to the general methods previously described.'
Most Loranthaceae, and Krameria, are woody
perennials and represent exceptions to the ob-
servation (Stebbins, 1938) that woody plants
generally have small chromosomes. Stebbins sug-
gested that mechanical difficulties might arise
during mitosis if long chromosomes had to divide
in the relatively short transverse dimensions of
the cambial initials.
Chromosomal aberrations-Chromosomal aber-
rations are detected occasionally in Phoradendron.
The most common are anaphase bridges which are
sometimes accompanied by acentric fragments,
suggesting that they are the result of crossing-over
in inversion heterozygotes. Micronuclei are de-
tected occasionally at interkinesis and telophase
II.
LITERATURE CITED
BALDWIN, J. T., JR., AND B. M. SPEESE. 1!l57. Phora-
dendron flauescens: chromosomes, seedlings, and hosts.
Amer. Jour. Bot. 44: 136-140.
BILLINGS, F. H. 1932. Microsporogenesis in Phoraden-
dron. Ann. Bot. 46: 979-992.
1933. Development of embryo-sac in Phora-
dendron. Ann. Bot. 47: 261-278.
Co VAS, G. 1949. Estudios cariologicos en anthofitas III.
Darwinians 9; 158-162.
- - - , AND B. SCHNACK. 1946. Numero de chromo-
somas en antofitas de la region de Cuya (Republica
Argentina). Rev. Argentina Agron. 13: 152-166.
4 Slides of Trilliuni and Tradescantia were supplied
through the courtesy of Dr. Sherwin Carlquist.
[Vol. 51
DOWDING, E. S. 1931. Floral morphology of Arceutho-
biurn arnericanurn. Bot. Gaz. 41: 42-51.
GILL, L. S. 1935. Arceuthobiurn in the United States.
Trans. Conn. Aead. Arts and Sci. 32: 111-245.
- - - , AND F. G. HAWKSWORTH. 1961. The mistletoes:
A literature review. Tech. Bull. No. 1242. U.S. Dept.
Agric., Forest Service, Washington, D.C.
KING, R. M. 1961. Meiotic chromosome numbers for
two species of the genus Psittacanthus (Loranthaceae).
Southw. Nat. 6: 48-49.
LAWRENCE, G. H. M. 1951. Taxonomy of vascular
plants. The Macmillan Co., New York.
MOORING, J. S. 1960. A cytogenetic study of Clarkia
unguiculata II. Supernumerary chromosomes. Amer,
Jour. Bot. 47: 847-854.
MUNTZING, A. 1958. A new category of chromosomes.
Proc. 10th Internatl. Congr. Genet., 453-467.
PISEK, A. 1924. Antherenentwicklung und meiotische
Teilung bei der Wacholdermistel [Arceuthobiurn
oxycedri (DC.) M. B.]: Antherenbau und Chromo-
somenzahlen von Loranihus europaeus Jacq. Sitzungs-
ber. Akad. Wiss. Wien, Math.-Nat. KI. Abt. I. 133:
1-15.
SHARP, L. W. 1943. Fundamentals of cytology. Me-
Graw-Hill, New York.
STEBBINS, G. L., JR. 1938. Cytological characteristics
associated with different growth habits in the dicotyle-
dons. Amer. Jour. Bot. 25: 189-]9S.
1950. Variation and evolution in plants.
Columbia University Press, New York.
SWANSON, C. P. 1957. Cytology and cytogenetics.
Prentice-Hall, Inc., Englewood Cliffs, New Jersey.
TRELEASE, W. 1916. The genus Phoradendron: a
monographic revision. University of Illinois Press,
Urbana, Illinois.
TURNER, B. L. 1!l58. Chromosome numbers in the
genus Krameria: evidence for familial status. Rhodoru
60: 101-106.
WIENS, D. 1964. Revision of the acataphyllous species
of Phonulendron: Brittonia (in press).