Safflower in California - University of California Cooperative Extension

Safflower in California 

The Paulden F. Knowles personal history of plant 

exploration and research on evolution, genetics, and breeding

Safflower in California 

The Paulden F. Knowles personal history 

of plant exploration and research 

on evolution, genetics, and breeding 

edited by 

Patrick E. McGuire 

Ardeshir B. Damania 

Calvin O. Qualset

Citation: 

McGuire, P.E., A.B. Damania, and C.O. Qualset (eds.) 2012. Safflower in California. The Paulden F. Knowles 

personal history of plant exploration and research on evolution, genetics, and breeding. Agronomy 

Progress Report No. 313, Dept. of Plant Sciences. University of California. Davis CA USA. 

Cover photographs: P.E. McGuire 

Preface photograph: Photographer unknown, UC Davis Dept. of Plant Sciences collection. 

Text photographs: P.F. Knowles

Contents 

v Editors’ summary: P.F. Knowles’ genetic resources legacy 

ix Preface 

xi Foreword 

xii Dedication 

1 Introduction 

1 Early history in California 

2 Commercial establishment in California 

3 Expanding the germplasm base 

7 Distribution of cultivated safflower 

10 Preservation of germplasm 

10 Germplasm evaluation 

12 Species relationships in Carthamus 

15 Fatty acid composition of safflower oil 

16 Male sterility 

17 Miscellaneous traits 

19 References 

Tables 

3 Table 1. Itinerary of 1958 collection trip. 

4 Table 2. Herbaria visited en route in 1958. 

6 Table 3. Itinerary of 1975 collection trip. 

7 Table 4. Local names for safflower. 

11 Table 5. Selections of safflower with resistance to four races of Fusarium wilt and registered with Crop 

Science Society of America. 

11 Table 6. Summary of number and origin of Verticillium wilt-resistant introductions. 

18 Table 7. Association of flower colors with chromatography colors. 

Figures 

5 Figure 1. Route map of 1964–65 trip covering 31,000 miles. 

14 Figure 2. Genome evolution in genus Carthamus. 

iii

Photographs 

x Paul Knowles, UC Davis 

4 Two-row planter used for safflower in India in 1965. 

8 Minor use of safflower as a vegetable in 1965. 

9 Transporting harvested safflower in India in 1965. 

13 Threshing and winnowing safflower in India in 1965. 

16 Processing safflower for oil in Egypt in 1958. 

Appendices 

23 Appendix 1 Table 1. Count of accessions of cultivated safflower (Carthamus tinctorius) collected in 1958. 

25 Appendix 1 Table 2. Count of accessions of wild species of safflower (genus Carthamus) collected in 1958. 

26 Appendix 1 Table 3. Count of accessions of cultivated safflower (Carthamus tinctorius) collected in 

1964–65. 

27 Appendix 1 Table 4. Accessions of wild species of safflower (genus Carthamus) collected in 1964–65. 

28 Appendix 1 Table 5. Accessions of Carduncellus species collected in 1964–65. 

29 Appendix 1 Table 6. Safflower germplasm selected in China in 1988. 

33 Appendix 2. Publications of P.F. Knowles on oilseed crops. 

41 Appendix 3. List by species, year, and PI number of 1175 safflower accessions associated with P.F. Knowles 

in the collection at the USDA-ARS-RPIS, Pullman WA USA. 

iv

Editors’ summary: 

P.F. Knowles’ genetic resources legacy 

At the start of Paul Knowles’ career, safflower was only incidentally grown in California. By 

virtue of his work, the crop was introduced and improved and production and distribution became a California 

industry. Central to this progress was germplasm acquisition, evaluation, conservation, and utilization carried out 

with Knowles’ vision and leadership. His achievements were far reaching. Today, when crop genetic resources are 

paradoxically both increasingly recognized as highly valuable and yet confronted with ever greater risk of being 

lost, his story has valuable lessons. 

• Accessible, wide-ranging genetic diversity is essential for the genetic improvement of a crop. 

Knowles recognized early on that safflower breeding programs in the US had a narrow genetic base and 

this lack of germplasm was restricting progress. 

• Accessibility of genetic diversity requires international exchange and collaboration. 

Knowles was supported, directly and indirectly, in his efforts by his institution, several US agencies (e.g., 

USDA, USAID, US Dept. of State), the UN Food and Agriculture Organization, and numerous academic 

and national researchers and institutions and private growers and sellers in the many countries in which he 

traveled. He initiated and facilitated exchanges of germplasm, information, and scholars. 

• Collection of germplasm should be made across all environments and geographic regions in which the 

target species or crop grows. 

The mutations he reports and worked with for safflower fatty acid composition arose in very small populations 

in widely dispersed and unpredictable sites (India, Iran, Israel, Portugal, and Russia) and would never 

have been found without access to wide-ranging germplasm collections. 

• Useful genetic diversity can be found in landraces of the crop from regions where it has a long history of 

cultivation and in wild species closely related to the crop species. 

He reports useful genes and traits for cultivated safflower from landraces (e.g., the fatty acid composition 

mutations, flower color variants, growth habit, disease resistances, and male sterility) and from wild species 

(disease resistances). 

v

• Indigenous germplasm is being rapidly replaced with introductions and newly developed varieties in 

many regions and environments, it is critical that material continues to be collected for ex situ conservation 

from these areas. 

He recognized early in his collecting career that safflower landraces that had been grown for centuries in 

some countries were being replaced by crops and cultivars from developed nations. 

• Research on species relationships and distribution, ecology, ethnobotany, cytogenetics, and basic biology 

are crucial for efficient and directed utilization of landraces and wild species for improvement of 

related crops. 

Over his career, his graduate students, postdocs, and research colleagues compiled a broad record of research 

efforts on safflower that encompassed basic agronomy, pathology, cytogenetics, genetics, biochemistry, 

and biogeography. The productivity of this research effort depended in a large way on the safflower 

genetic resources collection (the World Collection) to which he was a major contributor. At the same time, 

these research efforts enhanced the value of the World Collection as the resulting data became part of the 

information associated with the collection. 

• Evaluation and characterization of genetic resources (landraces and breeding lines) under diverse environments 

and stress conditions are essential for identifying characters useful for crop improvement. 

His work in California with the World Collection and screening for disease resistances and adaptive traits 

and the work with collaborators in other countries and environments exemplify the value of the evaluation 

and characterization process for ultimately successful germplasm utilization. 

• Novel genetic diversity can lead directly to successful cultivar development. 

Safflower cultivar ‘UC-1’, released by Knowles in 1968, was essentially the first commercial variety of a new 

type of safflower oil, a new crop. Its oil properties (high oleic acid) differed in fatty acid composition from 

that of traditional safflower oil (high linoleic acid) as a result of a mutation discovered in a safflower accession 

from India. The oil of UC-1 was chemically similar to olive oil. The identification of the novel oil type 

was made possible only by screening diverse collections of safflower germplasm for variation in fatty acid 

properties. The Indian accession was crossed in Knowles’ breeding program to a Nebraska cultivar (N-10) 

in 1957 to initiate the incorporation of the novel oil type into an agronomically acceptable background. By 

1967, after a backcrossing breeding program, certified seed of the new cultivar was available and it was the 

foundation for California production for several years and for commercial breeding programs thereafter. 

• Crop introduction is facilitated by agronomic research and outreach and extension to cultivate growers, 

producers, and markets for production. 

His work in this area early in his career laid the foundation for all his subsequent work. The rising success 

of safflower as a California crop gave impetus and support for his subsequent breeding, evolution, and 

geographic interests in the crop and its wild related species. 

• Plant ‘breeding’ in the broadest sense of plant selection, use, and improvement is a story of human curiosity, 

need, innovation, and persistence. 

On his travels, he didn’t interact only with breeders and seed dealers, he sought out farmers, market sellers, 

and users of the crops in which he was interested. This report on safflower touches on some of the many 

anecdotal presentations of the ways of growing, harvesting, processing, and using (in art, medicine, industry, 

and cuisine). The similarities and contrasts that he observed from one country and tradition to another 

in all these steps with a common plant are enlightening and a testimony to his own sense of curiosity and 

the apparent value he placed on each person’s contribution, testimony, and information. 

• Serendipity plays a significant role in revealing the utility of germplasm collections. 

Two examples from Knowles’ work illustrate this: 

1. Lettuce is known commonly as a leafy vegetable, and more distantly in time, it was cultivated for its 

stem, but on both his 1958 and 1964–1965 collection trips, he observed the use of a primitive lettuce 

vi

in Egypt as an oil crop in areas where safflower was also cultivated. Its seeds were being collected and 

pressed, in a manner similar to how safflower oil was obtained. Seed derived from his lettuce collections 

is still available through USDA National Plant Germplasm System (NPGS), e.g., PI 250020. 

2. Among the many other non-Carthamus taxa that he collected during his 1958 trip were four ‘forage 

grass’ accessions that he labeled ‘Agropyron spp.’ in his report (Knowles 1959), but only ‘grass’ or 

‘annual grass’ in his notebook. These were accessioned by the USDA NPGS and identified to species. 

Three of them turned out to be Brachypodium distachyon: PI 253334 was collected in Morocco and 

PI 254867 and PI 254868 were collected in Iraq. All three were screened in the last decade as part 

of a large-scale effort1 at understanding the geographic, morphological, and genetic diversity in this 

species in preparation for presenting it as a new model plant system for genomic research. An inbred 

line (denoted Bd21) derived from Knowles’ PI 254867 accession (seed he collected in Iraq as K 1202, 

‘4 km. from Salahudin, 4"-6" tall, ripe’, coming from Mosul) became adopted, primarily because it was 

diploid and because of its facility for transformation, as the ‘canonical genotype’ for B. distachyon serving 

as the source of DNA and RNA for both the whole genome and EST sequencing projects by the 

US Dept. of Energy-Joint Genome Institute Community Sequencing Project (DOE-JGI). The full genome 

sequence for B. distachyon from Bd21 was published in 2010. 2 In 1958 Knowles opportunistically 

collected those seeds and deposited them in the NPGS. Some 40 years later, a testament to NPGS 

maintenance and regeneration practices, the accessions were thus available for uses and technologies 

completely unknown in 1958. 

Paul Knowles’ work finished with the decade of the 1980s. At the time of his death in 1990, work was underway that 

would culminate with the opening for signature in 1992 of the Convention on Biological Diversity and its subsequent 

entry into force at the end of 1993. It is an important question whether he could have done his work in the 

international germplasm access and exchange environment that exists post-CBD. Certainly under the CBD, there 

is nothing in theory that would prevent his accomplishments, but in practice the many bilateral agreements for 

exchange of germplasm necessary today and the difficulty in obtaining these (as exemplified by the records of the 

past 20 years) make it highly unlikely that the current state of safflower knowledge and productivity would have 

been possible. The International Treaty for Plant Genetic Resources for Food and Agriculture and its Multilateral 

System for genetic resources access that emerged in the early 2000s would not have helped Knowles’ safflower 

work either. Safflower is not one of the crops covered under the Treaty. 

— Patrick E. McGuire, Ardeshir B. Damania, and Calvin O. Qualset 

Department of Plant Sciences, University of California, Davis CA USA 

1 Vogel JP, DF Garvin, OM Leong, and DM Hayden. 2006. Agrobacterium-mediated transformation and inbred line development 

in the model grass Brachypodium distachyon. Plant Cell, Tissue and Organ Culture 84:199–211; 

Garvin DF, Y-Q Gu, R Hasterok, SP Hazen, G Jenkins, TC Mockler, LAJ Mur, and JP Vogel. 2008. Development of genetic and 

genomic research resources for Brachypodium distachyon, a new model system for grass crop research. Crop Sci 48(S1):S69–S84. 

2 The International Brachypodium Initiative. 2010. Genome sequencing and analysis of the model grass Brachypodium distachyon. 

Nature 463:763–768. 

vii

Preface 

The history of safflower in California is a classic example of the importance of acquisition and use 

of genetic resources in the adaptation and spread of a crop in an agroecosystem. Paulden F. Knowles (b. April 18, 

1919, d. February 7, 1990) played a key role in the establishment of safflower as a California crop. Shortly before his 

death he began a personal account of that history. He undertook this at the request of the University of California 

Genetic Resources Conservation Program (GRCP), which intended its publication as number 14 of its numbered 

report series. Unfortunately, the report was never completed and published before GRCP was closed in 2008. 

Because safflower is such a good model for crop development via germplasm acquisition, conservation, and 

utilization and because Knowles’ vision and achievements were so far reaching and productive, we think his story 

is still valuable even some twenty years later. Accordingly we returned to Paulden Knowles’ hand-written first draft 

and his notes about plans for the report and produced this current document. 

The itineraries and results of his collecting trips are even more impressive today, given the policy constraints at 

both international and national levels and the subsequent changes in accessibility of genetic resources over the past 

20 years on the one hand, and the political and armed conflicts and environmental changes in the regions in which 

he collected on the other. These collections can never be replicated. 

His collections were not a one-directional transfer of germplasm and information. Along the way he documented 

and shared usage and cultural information about the materials he collected. His travels forged the links of a 

network of researchers involved with safflower. One manifestation of this network were the graduate students from 

many countries who came through his program at the University of California, Davis. Another was the initiation of 

international safflower conferences with broad participation by researchers from countries in which safflower is an 

important crop. 

There had been US national safflower conferences beginning in the early 1960s. Dr. Knowles was instrumental 

in organizing the Third Safflower Research Conference, held in 1969 at the University of California, Davis CA and 

edited the proceedings of that with M.D. Miller. By the late 1970s, however, the importance of a broader perspective 

on safflower research and utilization led to planning for a conference on an international scale. The first one 

was held in Davis CA in 1981, with Dr. Knowles as the organizer and subsequently editor of that proceedings volume. 

At that conference, he was one of two award recipients; he was recognized for his service as ‘scientist, teacher 

and administrator’. The other was to Carl Claassen as ‘innovative oilseed scientist and agribusiness leader’. The second 

international safflower conference was eight years later in 1989 in India. At this conference, Dr. Knowles was 

the keynote speaker in recognition of his contributions, being introduced as ‘world’s renowned scientist and father 

of safflower’. Since then the conferences have continued on a four-year cycle, moving from India to China, Italy, the 

US again, Turkey, Australia, and India again, in January of this current year, with the 8th conference. 

ix

His work enriched the germplasm holdings of the US National Genetic Resources Program (NGRP), which 

has been the ultimate repository for his material. In addition to the germplasm, the system has characterization 

and evaluation data on the accessions from Dr. Knowles’ work and that of many other researchers. Safflower was 

not the only target for Dr. Knowles either: his collection activity and research also dealt with other existing and 

potential oil crops such as flax, sunflower, soybean, brassicas, crambe, sesame, cuphea, and castor, as well as forage 

grasses and herbs. 

We note two volumes that provide comprehensive information of safflower as a crop and testimony to Dr. 

Knowles contribution. One is Oil Crops of the World (1989, McGraw-Hill Publishing Co., New York NY USA, 

xviii+551 pages), edited by G. Röbbelen, R.K. Downey, and A. Ashri, to which Dr. Knowles contributed the chapter 

on safflower and the chapter on genetics and breeding of oil crops. The book was dedicated to Paul Knowles. 

The second is Safflower (1996, AOCS Press, Urbana IL USA. xiii+592 pages) by J.R. Smith. This work, from the 

perspective of Smith’s career in the safflower industry, provides a throrough history of safflower as a crop and documentation 

of Knowles’ contributions. 

Note on the text. The text is based on Dr. Knowles’ long-hand manuscript supplemented by notes and his outline 

of the document. Some updates are provided as footnotes, but no effort has been made to create a review of the 

advances in safflower research and crop development that have taken place since he drafted the manuscript. The 

tables in the text and in Appendix 1 were planned by him. Illustrations have been added from his extensive unpublished 

photograph and slide collection. 

In addition, we have compiled in Appendix 2 a bibliography of his published work on oil crops and in Appendix 

3 a listing of the accessions associated with Dr. Knowles of safflower and wild Carthamus-species relatives maintained 

at the USDA-ARS-Regional Plant Introduction Station, Pullman WA USA. The actual number of introductions 

for which Knowles was responsible of safflower and related species in Carthamus and other genera related to 

Carthamus is greater than these 1175 accessions. Some accessions of Carthamus were no longer viable and maintained 

at the RPIS by 1995 and thus were not in this listing. In addition, he was responsible for collections of seed 

from species of other genera related to Carthamus, but not listed in Appendix 3 (for example, see Appendix 1 Table 

5). As we note in the Editors’ Summary, 

he also collected seed from species of 

plant families other than Asteraceae and 

accessions of many of these are maintained 

in the USDA NGRP. 

We acknowledge contributions to Dr. 

Knowles in 1989 for the preparation of his 

manuscript by Amram Ashri (review), 

A. Lee Urie (summary of USDA activity 

at Davis), and Richard C. Johnson (summary 

of Knowles accessions in GRIN) 

and to us in 1995 by Raymond L. Clark 

(information on Knowles accessions in 

GRIN) for the preparation of this report. 

— Patrick E. McGuire, Ardeshir B. 

Damania, and Calvin O. Qualset 

Department of Plant Sciences, University 

of California, Davis CA USA 

Paul Knowles at his desk at the University of California, Davis. 

UC Dept. of Plant Sciences Collection. 

x

Foreword 

The story of Paulden F. Knowles and safflower in California exemplifies the trials and tribulations and 

hopes involved in the introduction of a new crop far from its original habitat. It shows the many vital interactions 

between farmers, processors, and research and extension workers which are crucial to success of the crop. Knowles 

has shown the way through the development of safflower in California, which can serve as a model case study on 

the future development of new crops, a topic which is of great significance these days in both the developed, but 

gene-poor countries of the north, as well as the developing, but gene-rich countries of the south. 

Knowles was immensely successful in introducing a balanced research program which involved the collaboration 

of the USDA, the private industry with commercial interests, and the University of California’s efforts. This 

he could do because of his scientific stature, motivation, ability to foresee long-term needs of the industry, and 

personal attributes, as was the case with his collection of germplasm, evaluation, wild species research, and genetic 

manipulations of oil quality. 

Last but not the least, Knowles was a research leader, a scholar, an educator who attracted, taught, and inspired 

many of his students from all parts of the world. Today several of his students are carrying on the traditions of 

dedicated research and education imparted to them by the personal example of Knowles. 

— Amram Ashri, 1991 

Hebrew University of Jerusalem, Israel 

xi

Dedication 

This report is dedicated to my graduate students who worked on safflower. Without their enthusiasm, 

dedication to research, and imagination, this report would not be possible. I salute them. Their names and 

present locations 1 are given below. 

— Paulden F. Knowles, 1989 

1 Editors’ note: At time of Knowles’ draft, 1989. 

Amram Ashri, Israel 

Yousef Attieh, Jordan 

Sheldon Bartholomew, deceased 

Harold E. Bockelman, Idaho USA 

Jirair Carapetian, Iran 

John Dillé, South Carolina USA 

Ali Estilai, California USA 

Wesley W. Ebert 2 , California USA 

José Fernández Martínez, Spain 

Shama Futehally, Pakistan 

August Hartman, California USA 

Bryan Harvey, Canada 

2 Editor’s note: Deceased. 

xii 

Arthur B. Hill, California USA 

Thomas C. Heaton, California USA 

Bruce Imrie, Australia 

M. Osman Khidir, Iraq 

Sheldon Ladd 2 , Oregon USA 

Ricardo León, Mexico 

Osman Mutwakil, Sudan 

G.V. Ramanamurthy, India 

El Saeed El Saeed, Sudan 

Stanley S. Schank 2 , Florida USA 

Steven R. Temple, California USA 

Demetrius Yermanos, deceased

Introduction 

In October of 1947 I joined the staff of what was then called the Division of Agronomy of the University 

of California. My assignment was oilseed crops, primarily flax, which at that time was grown commercially in 

the Imperial Valley, the Westside of the San Joaquin Valley, and San Mateo County. While flax was given emphasis, 

an effort was made to assemble seeds of other oilseed crops. 

A fruitful source of seeds was the Chemurgy Project at the University of Nebraska at Lincoln. From Dr. C.E. 

Claassen, in charge of evaluations of several oil crops, seed was obtained of cultivars of safflower, castor, and 

sesame. Safflower (Carthamus tinctorius L.) was sown in late January, 1948, and other crops in the spring. Most 

promising of those crops was safflower, where in test plots Nebraska materials yielded 3011 lbs/ac (3372 kg/ha). 

In subsequent years safflower became the primary crop in my research program. This is an account of my 

involvement with that crop up to and beyond retirement in 1983. I am grateful both to graduate students, who did 

much of the work, and to members of other departments. Emphasis has been given to germplasm collection and 

evaluation. The account also includes results of research that developed from studies of introduced germplasm. 

Early history in California 

The University of California’s evaluations of safflower germplasm began about the turn of the 20th 

century. For trials at the several UC field stations in the period 1899-1901, Shinn (1903) summarizes: 

Foothill Station (near Jackson): The well-known safflower, Carthamus tinctorius (No. 1345 of the Inventory), was 

grown in this region years ago, and occasionally appears in old gardens. Since tested on the substation, it proves very 

easy of growth on both soils, very drought-enduring. Does not compare with the Russian sunflowers as a yielder of 

seeds for oil or chicken feed. 

Southern Coast Range Station (near Paso Robles): Safflower.—No. 1343, sown December 4th, grew three feet high, 

bloomed May 30th, and kept green until autumn. 

Southern California Station (Chino Valley): Seeds of this well-known dye and oil plant (Carthamus tinctorius), 

No. 1345 of the Inventory, were sown on both tracts. On the dry land (home tract) it grew 3 feet high and produced 

1

some seed; on the moist land the crop was very large, the plants being 4 feet high. It produced seeds at the rate of 

5,500 pounds per acre. The drought resistance of this plant appeared considerably greater at Paso Robles than at this 

substation. It thrives over a large part of California, and is of very easy culture. 

The first introductions of safflower to California were probably made much earlier when immigrants from the 

Mediterranean area brought with them seed for planting in their gardens. In the countries of origin of these immigrants, 

safflower had been grown for centuries for the red flowers which were used to color foods. They were a 

substitute for saffron, though they have a bland taste. 

The first large-scale testing program in the US was directed by Frank Rabak of the US Department of Agriculture 

(Rabak 1935). This led to small commercial plantings in the northern Great Plains and California. The low 

yields and low oil contents of introductions at that time, most of them from India, discouraged interest by oilseed 

processing companies. 

Commercial establishment in California 

The success of the initial nursery plantings in 1947–48 led to larger planting in 1948–49. The average 

yield of eight varieties and selections developed by the University of Nebraska was 2753 lb/ac (3083 kg/ha) 

(Knowles 1949b). Such yields and the good performance of safflower in small semi-commercial plantings totalling 

less than 100 acres provided the information that led to extensive commercial plantings of over 25,000 acres 

in 1949–50. Another stimulating factor was mandatory cutbacks in California’s cotton acreage, which led also to 

commercial plantings of castor. 

Two companies contracted acreages in the 1949–50 season, Pacific Vegetable Oil Corporation in San Francisco 

and Oilseed Products Company in Fresno. If average yields are considered, the crop was a disaster. The reasons 

were (Knowles and Davis 1951): 

• “A few growers put safflower on poor ground thinking that safflower would perform better than other crops 

under such circumstances. 

• “The variety N-852 proved to be very susceptible to root rot, and the disease was aggravated by untimely 

irrigations. 

• “Many disappointments with safflower were a consequence of late dates of seeding. 

• “A few plantings made early in the fall suffered from frost damage. 

• “Some loss from shattering occurred when the crop was allowed to stand for some time after it was ripe. 

• “Weeds and volunteer grain in some cases seriously reduced yields.” 

On the other hand, there were some excellent yields, one field on the west side of Fresno County gave over 

4800 lb/ac (5380 kg/ha). A field near Meridian in the Sacramento Valley gave about 3700 lb/ac (4140 kg/ha). 

The few successes outweighed the disasters. Several farmers elected to grow safflower in 1950–51, with only Pacific 

Vegetable Oil Company contracting acreages. 

Other factors were involved in the successful establishment of safflower, the most important being: 

1. An expansion in the information base: As expected, farmers learned quickly from their successes and 

failures and experience of their neighbors. The University of California at Davis and at the West Side Field 

Station, the Agricultural Extension Service through several farm advisors in the Central Valley, Imperial 

Valley, and Palo Verde Valley, and the USDA at the Cotton Research Station near Shafter, together provided 

a great deal of information (Knowles and Davis 1951). In addition, the Pacific Vegetable Oil Corporation 

conducted several nursery trials. 

2. Contracted acreages: The Pacific Vegetable Oil Corporation established contracts with growers which 

guaranteed both a market for the crop and a price tied to the selling price of the oil and meal but not below 

a stated minimum price. 

3. An advisory service provided by the Corporation which farmers could contact at any time before, during, 

and after planting. 

2

4. Promotion: The Pacific Vegetable Oil Corporation held many meetings both in fields and indoors during 

the winter. At such meetings University of California personnel contributed information from their 

research on safflower. 

From this point on, there was annual safflower production in California. The fluctuations in acreage were 

strongly influenced by the relative market prices of safflower and wheat. 

A major factor in the early years that adversely affected the commercial development of safflower was disposal 

of the meal. While the oil was moving in the marketplace, the meal was not. The Pacific Vegetable Oil Corporation 

provided funds to enable the University of California to evaluate the meal as feed for poultry and for livestock. 

With good data on comparisons of safflower meal with other oilseed meals, large accumulations of the meals were 

no longer a problem. 

Expanding the germplasm base 

It was not many years after the commercial establishment of safflower that it became apparent that 

breeding programs were increasingly restricted by lack of germplasm, most of which stemmed back to materials 

from the University of Nebraska. It was also known that many countries in the Old World were substituting crops 

and varieties from developed nations for those that had been grown for hundreds of years. 

Collection trip in 1958 

The University of California and the USDA agreed to fund a joint collection trip, whereby the University paid my 

salary and the USDA paid all expenses. My journey began on March 5, 1958, and terminated in early November 

(see Table 1 for itinerary). 

Most of the travel was by air 

and railroad, with local travel 

by rented vehicles or vehicles 

loaned or charged to me by 

the US Agency for International 

Development (USAID) 

or research stations. Wherever 

I went I was hospitably received 

and generously treated. 

A detailed report of the collection 

trip was filed with the 

Department of Agronomy, 

UC Davis and the New Crops 

Research Branch, Crops Research 

Division, ARS, USDA 

(Knowles 1959). A total of 

420 accessions of cultivated 

safflower (Appendix 1 Table 

1), 431 accessions of wild 

Carthamus species (Appendix 

1 Table 2), and 29 accessions 

of species of other wild genera 

related to Carthamnus were 

collected. 

Table 1. Itinerary of 1958 collection trip. 

Date Location 

April 4–24, May 21 India (Delhi, Pusa, Hyderabad, Poona, Amritsar) 

April 25–May 20, 22–24 Pakistan (Lahore, Lyallpur, Rawalpindi, Peshawar, Karachi) 

May 25–29 Afghanistan (Kandahar, Lashkar Gah—southern areas) 

May 30–June 4 Iran (Tehran, Abadan, Ahvaz, Dezful—southern areas) 

June 5–10 Iraq (Baghdad, Hilla, desert west of Ramadi—southern areas) 

June 11–20 Egypt (Cairo, Alexandria, Kena, Luxor, Sohag, Assiut) 

June 21–24 Jordan (Amman, Deir Alla Station, Nablus, Jerusalem) 

June 25–28 Syria (Damascus, Aleppo) 

June 29–July 9 Iraq (Mosul, Salahudin, Sirsank, Kirkuk, Sulaimaniya—northern areas) 

July 10 31 Iran (Tehran, Isfahan, Tabriz, Mashad) 

August 1–10 Afghanistan (Kabul, Ghazni, Dasht-I-Nawar, Charikar—central areas) 

August 11 Iran (Tehran) 

August 12–24 Turkey (Ankara, Gokhoyuk, Amasya, Tokat, Yozgat, Eskişehir, Bursa, 

Balikesir, Izmir, Istanbul) 

August 25–31 Israel (Tel Aviv, Jerusalem, Beersheba, Tiberias, Nazareth, Safad, Acre, Haifa) 

September 1 France (Paris) 

September 2–9 Morocco (Rabat, Oujda, Melilla, Casablanca, Tadla, Bine el Quidane, Meknes) 

September 10–20, 25–26 Spain (Madrid, Cuenca, Teruel, Albacete, Alicante, Elche, Murcia, Granada, 

Malaga, Seville, Trujillo) 

September 21–24 Portugal (Lisbon, Sacavém, Oeiras) 

3

Herbaria. As plans were made for the collec- Table 2. Herbaria visited en route in 1958. 

tion trip it was soon apparent that there was Date Herbarium/location 

very little information on where to collect. To 

remedy this situation I examined herbarium 

specimens of Carthamus at the University of 

California in Berkeley; Stanford University; 

March 6–7 

March 10 

March 12–14 

US National Herbarium, Smithsonian Institution, Washington DC USA 

New York Botanical Garden, Bronx Park, New York NY USA 

Royal Botanic Garden, Edinburgh, SCOTLAND 

the California Academy of Sciences, Golden March 17–20 Royal Botanic Garden, Kew, Surrey, ENGLAND 

Gate Park, San Francisco; the State Depart- March 19 Linnaean Herbarium, Burlington House, London, ENGLAND 

ment of Agriculture, Sacramento; and the 

Rancho Santa Ana Botanic Garden, Claremont, 

California. In addition, en route to the 

collection areas, herbaria in several countries 

March 21 

March 24–25 

May 5 

British Museum, London, ENGLAND 

Museum Nationale d’Histoire Naturelle, Paris, FRANCE 

Gordon College, Rawalpindi, PAKISTAN 

were also visited (Table 2). 

June 5 Abu Ghraib Experiment Station, Abu Ghraib, IRAQ 

Those herbaria provided useful informa- June 9 University College of Arts and Science, Baghdad, IRAQ 

tion on locations where safflower had been June 16 University of Cairo, Cairo, EGYPT 

collected over a period of more than 100 

years. Where collections were dated it was 

possible to identify the best time of year to 

make collections. Brief notes and sometimes 

June 21 

August 13 

August 26 

Deir Alla Station, Ministry of Agriculture, JORDAN 

University of Ankara, Ankara, TURKEY 

Hebrew University, Jerusalem, ISRAEL 

photographs were taken of the specimens. September 3 Institut Scientifique Chérifien, Rabat, MOROCCO 

Such information for the cultivated species September 11 Jardín Botanico, Madrid, SPAIN 

and wild (weedy) species with 10, 11, 12, 22, September 22 Estaçäo Agronómica Nacional, Sacavém, PORTUGAL 

and 32 pairs of chromosomes were compiled 

along with information on materials loaned 

October 1–3 Jardin Botanique, Geneva, SWITZERLAND 

by European herbaria subsequent to the visits made during the collection trip. 

Collection trip in 1964–65 

On a sabbatic leave for a year, I again visited much of the area visited in 1958, but drove a mini-bus for about 31,000 

miles (and 32 flat tires), most of the distance covered on the trip. With my own car I was able to visit out-of-theway 

places. I was accompanied by my wife for the entire trip and by my son for the first six months. In India we had 

access to government rest houses which were inexpensive, but required that we carry our own bedroll and mosquito 

netting (Knowles 1965a). Our route and side-trips by air or sea are indicated in Figure 1. 

Time was taken to search for and obtain 

photos of systems of growing safflower. 

Village-level processing facilities have been 

described and photographed (Knowles 

1967). A total of 519 accessions of cultivated 

safflower (Appendix 1 Table 3), 136 accessions 

of wild Carthamus species (Appendix 

1 Table 4), and 36 accessions of Carduncullus 

species (Appendix 1 Table 5) were 

obtained. 

Collection trip in 1975 

It was realized that a few areas of the Near 

East had not been visited in a search for 

safflower. These were: Lebanon, western 

and eastern Turkey, and western Iran (see 

Table 3 for itinerary). Following a study of 

oilseed crops research needs in Egypt for 

the UN FAO in 1975, I extended my travels 

Two-row planter used for safflower. Hatta, Maharashtra State, India. 

Feb. 28, 1965. (Text and photos from Knowles 1965a.) 

4

Figure 1. Route map of 1964–65 trip covering 31,000 miles. 

5

to visit those areas. I was fortunate 

to visit Lebanon during a brief lull in 

the exchanges of rifle fire. 

In Lebanon, the Ford Foundation 

provided me with a car and 

driver. I rented a car for travel alone 

in western Turkey, and in eastern 

Turkey with the help of Dr. Enver 

Esendal, on the staff of Atatürk 

University in Erzurum, I was able to 

rent another car with a driver. Dr. 

Esendal was my guide for the entire 

journey. In western Iran, Mr. H.G. 

Khadivi, Managing Director of the 

Oilseed Research and Development 

Company in Tehran, provided me 

with his personal car and driver. 

The late N.A. Ghanavati, the leading 

oilseed breeder in Iran at that time 

based at the Seed and Plant Improvement 

Center, Varamin, was my 

guide and interpreter. I am indebted 

to these persons or agencies for their 

generous assistance. 

Germplasm from China, 1988 

Over a period of seven or eight 

years letters and seed samples were 

exchanged with Dr. Li Dajue at the 

Beijing Botanical Garden, Institute 

of Botany, Chinese Academy of Sciences, 

Xiangshan, Beijing. This led 

to a visit which my wife and I made 

to China in 1988. We bore all our 

own expenses except accommodation 

and travel expenses for me in 

Table 3. Itinerary of 1975 collection trip. 

Date Location 

Lebanon 

July 13 Arrive Beirut 

July 14 To the Bekaa Valley, north to Kfardan Research Station and south to the Tal Amara 

Research Station. Stayed at Zahle. 

July 15 Drove south along west side of Bekaa Valley to the end of the lake, then back 

through the center of the Valley and return to Zahle. The only wild species of safflower 

seen here was C. anatolicus. 

July 16 Drove north and east to Baalbek, then NW to Les Cedres, and return to Beirut via 

Becharra. 

Turkey 

July 17–22 Beirut to Ankara, Turkey. Visited research stations and collected in the Ankara area. 

July 22 To Izmir in western Turkey. 

July 23–24 Visited Agricultural Research and Introduction Center, P.K. 9, Menemen, Ege 

University, and a University station west of Izmir. 

July 25–26 Drove south to Bodrum via Efes (Ephesus), then via Mugla, Köyceğiz, and Dalaman 

to Feth’ye on the Mediterranean Sea. 

July 27 Returned to Izmir by an inland route via Kemer, Altinyayla, Gölhisar, Acıpayam, 

Denizli, and Aydin.. 

July 29–30 To Erzurum in eastern Turkey. Visit to Atatürk University and area of Cat. 

July 31– Traveled east via Paşinler, Horasan, Tahir, and Eleşkirt, then south via Tutak, 

Aug. 3 Patnos, Kocapinar, Erciş, and Timar to Van (7/31); then to Mardin via Tatvan, 

Bitlis, Baykan, Kurtalan, Besiri, Gercüş, Midyat, and Savur (8/1); then to Bingöl 

via Diyarbakir and Elazig (8/2); and return to Erzurum via Mus, Varto, Hınıs, and 

Paşinler (8/3). 

Western Iran 

Aug. 5–6 To Rezaiyeh in NW Iran via Tabriz. 

Aug. 7–9 In Rezaiyeh area 

Aug. 10–13 To Bukan via Mahabad and Mujandoah (8/10); to Kermanshah via Sarandaj 

(8/11); to Khorramabad via Shahabad, and Malavi, then to Hamadan (8/12); and 

to Tehran via the main highway (8/13). 

Aug. 14–16 Visiting research stations near Tehran. 

Aug. 17 Flew to Rome enroute to Spain. 

China. We traveled both by plane and by train. Safflower nurseries were visited at: the Beijing Botanical Garden; 

the Qinghai Academy of Forestry and Agriculture, Xining, Qinghai Province; Institute of Industrial Crops, Chinese 

Academy of Sciences and Technology, Urumqi, Xinjiang Autonomous Region; and the Station of Agricultural 

Technology and Extension, Wengniute Qi, Chifeng City, Inner Mongolia. 

Dr. Li had grown a nursery of Chinese germplasm at Beijing, all of it under a plastic cover to prevent both 

access of insects which would cause cross-pollination and exposure to rain which would lead to serious disease 

attacks. He shared seed from that planting and it was shipped (93 accessions along with 10 accessions collected 

by me during this trip, see Appendix 1 Table 6) via the American Embassy to the USDA Germplasm Resources 

Laboratory, Beltsville MD. There the collection was accessioned and sent to the USDA Regional Plant Introduction 

Station in Pullman WA for seed increase and characterization. 

At my request the USDA had supplied Dr. Li with seed of all safflower entries in the National Plant Germplasm 

System. A single-replicate nursery was grown at the Station of Wengniute Qi, near Chifeng, Inner Mongolia. With 

the assistance of Dr. Li and others, superior plants or rows that were spineless and orange or red flowered were 

identified. These plants, like other germplasm that I had sent to Dr. Li, will be used in breeding programs at several 

stations. 

6

Distribution of cultivated safflower 

This discussion consolidates information gained on collection trips in 1958, 1964–65, and 1975 (see 

Appendix 1 Tables 1, 3, and 6), including some information obtained in subsequent years. 

Safflower was found in every country that was visited from Turkey to Bangladesh, except for Lebanon. It was 

not distributed uniformly. It is grown in the Murcia-Elche area of Spain where saffron is grown, and a few plants 

were seen in 1965 near Tavira in southern Portugal. No safflower was seen in southeastern Europe. I have no evidence 

of cultivated safflower being indigenous to any area from Alexandria in Egypt to Rabat in Morocco, except 

on a visit to the Kufra Oasis in Libya in 1969 where I did see a small planting of a spineless, red-flowered type 

which presumably was grown for the flowers. Local names for safflower varied greatly. Those which I encountered 

and those derived from the literature (Ashri 1957) are given in Table 4. 

Safflower Centers 

From information available after the collection trips there appeared to be seven centers of safflower production in 

the Old World (Knowles 1969). These are described below. 

India—Pakistan Center. India leads 

all countries in safflower production. 

Most of it is grown in the south-central 

area in what is called the Deccan. 

The safflower of Pakistan is closely 

related to that in India. The remarkable 

feature of Indian safflower, 

except for recently developed types, 

is its uniformity; with few exceptions 

plants are spiny, orange flowered, 

bushy, and early (Knowles 1969). 

On average, for collections in 1965, 

99.3% were spiny and 98.9% were 

orange flowered. There is strong 

selection pressure against spineless 

types by birds and livestock. Bushy 

types would be favored because 

thin stands are common. It is hardly 

possible that orange flowers are a 

hold-over from a period over 150 

years ago when orange and red flowers 

were a source of carthamin, an 

important dye of commerce. If so, 

red-flowered types should have been 

more frequent. It is possible that the 

original introduction to the Deccan 

was an orange-flowered, spiny type. 

At the present time it is grown for 

oil, much of it extracted using simple 

village-level equipment. The cake 

residue is used as a livestock feed. In 

many areas safflower seedlings up to 

20 cm in height are used as a cooked 

vegetable. Sometimes seedlings are 

Table 4. Local names for safflower. 

Country and area Local name 

China hong hua (red flower) 

India Bihar State kusumba 

Hyderabad area kusuma 

Other areas kusum, karadai 

Kashmir hubulkhurtum 

Sanskrit cusumbha*; kamalotarra* (supreme beauty) 

Pakistan khurtum 

Afghanistan Kabul muswar or maswarah 

Herat kajireh 

Ghazni kariza 

Iran Isfahan kafsha, kafshe, or kosheh 

Tehran kafsha 

Tabriz zafaran-golu (Turkish) 

Rezaiyeh kouchan gule 

South of Rezaiyeh in Kurdish area kah’li 

Ghom golbar aftab 

Meshed kajireh or golzardu 

Shiraz khasdonah or laba torbak 

Saveh kajena goli or khardam 

Shahabad brarta 

Malavi kharkhool 

Iraq, Jordan, Syria, and Egypt (Arabic) qurtum, gurtum, osfur, or asper 

Turkey aspir or dikken (thistle) 

Sudan Kosheh* 

Ethiopia ssuff* 

Israel (Hebrew) khariah* 

* from Ashri (1957) 

7

“nipped” back to provide more plant as vegetable; it is believed by many that this practice will increase branching 

and yields. 

Middle East. Much of the safflower in the Middle East is grown for the flowers which are harvested when fresh 

and then dried. They may be found in bazaars of larger towns or cities, often in stores that sell dried saffron. The 

dried flowers, like dried saffron, are used to color foods such as rice, breads, and soups. In contrast to saffron, the 

flowers of safflower have a bland taste. In many instances where saffron is pulverized or powdered before being 

offered for sale, it is adulterated with pulverized or powdered flowers of safflower. As might be expected, these safflower 

plants are mostly spineless and red or orange flowered. 

Egypt. The flower colors of Egyptian safflower today are similar to that grown in 1600 BC, when the individual 

florets were sewn crosswise on long strips of fabric or papyrus to provide garlands for mummies. Present-day seeds 

are similar in shape and size to those used in Roman times when they probably served as a source of oil. Dried 

flowers are for sale in cites and larger towns. Some seed is sold as feed for larger birds. The major use of safflower 

seed is as a source of oil which is extracted in villages and towns of Upper Egypt with primitive equipment. In 1965 

it was apparent that the safflower of Upper Egypt was being strongly modified by spiny, yellow-flowered Nubian 

types which, following the completion of the Aswan High Dam, had been carried along by the people migrating 

from Nubia to Komombo, about 15 miles north of Aswan. Many tests of American varieties in the Nile Valley will 

have influenced the characteristics of Egyptian safflower. 

Sudan. The center of safflower production was in northern Nubia, the area along the Nile River in northern Sudan 

and southern Egypt, an area that is now largely covered by Lake Nasser behind the Aswan High Dam. There have 

been a limited number of Sudanese materials introduced into the US. Most have been yellow flowered, some have 

been orange flowered, and all have been spined, some strongly so. Two introductions from Sudan by the University 

of Nebraska showed commercial promise. Of these, N-852 became the commercial variety that was first used in 

commercial production in Nebraska, Colorado, and California. It was replaced by N-10, a selection from N-852. All 

varieties presently grown commercially stem back to N-852. Undoubtedly, prior to the construction of the Aswan 

High Dam, there was some migration of Sudan germplasm down the Nile River into Egypt. 

Ethiopia. Vavilov (1951) had proposed Ethiopia as a primary center in the evolution of safflower. This is hard to 

accept because there are no wild species related to cultivated safflower in Ethiopia—the only wild species reported 

to be from Ethiopia has 32 pairs of chromosomes (Khidir and Knowles 1970a). All introductions from Ethiopia 

that I have seen are similar; they 

are tall, spiny, many branched, red 

flowered, and small headed. The 

Ethiopian Center concept requires 

more study. 

Europe. At one time apparently, 

when safflower was grown as a 

source of carthamin, it was widely 

grown in Europe. Some relic 

populations undoubtedly still exist 

in warmer, drier areas, and several 

herbaria maintain seed stocks in 

small quantities. Flower colors are 

variable, and plants may be spiny or 

spineless. Field-scale plantings of 

up to one or two hectares in size are 

grown in the Murcia-Elche area of 

Spain where the prevailing type is 

red flowered, heads are intermediate 

in size, leaves are spineless to weakly 

A minor use of safflower in India (and Egypt) was as a vegetable. Seedlings 

(left) are harvested, removing roots and part of the stem (center), chopping 

them up (right), and cooking them like spinach , either by boiling or by 

cooking in hot oil. (Text and photo from Knowles 1969a.) 

8

spined, and there is an intermediate level of branching. The plantings of safflower are in the areas of Spain devoted 

to saffron production, suggesting that dried flowers of safflower may be used to adulterate saffron. 

Far East. At the time that the introductions from the collection trips were studied, there were very few introductions 

from the Far East, so that center must be considered as provisional—it will need major revision. 

At the First International Safflower Conference held at the University of California, Davis, June 12–16, 1981, Wu 

Ying-Siang and Li Dajue reported that 90% of the provinces in China grow safflower, mostly for its flowers (Wu 

and Li 1981). Li (1989) reported on the status of safflower research in China at the Second International Safflower 

Conference held in Hyderabad, India, January 9–11, 1989. The flowers are widely used solely or in admixture for 

medicinal purposes. Its primary benefits are reported to be to the circulatory system, particularly in the treatment 

of heart problems. It is also used for muscular strain, hypodermic congestion caused by bedsores, fracture of the 

bones, sprains, edema, abdominal distention, and problems of menstruation in women. About 1700 tons of dried 

flowers are processed each year. A few small factories extract the yellow flower color and red flower color (carthamin) 

from the orange- and red-flowered types, respectively. The former is being tested as a food dye to substitute 

for synthetic yellow colors which are in widespread use. The red color is used as an ingredient in medicines 

and probably as a food coloring. Safflower is reported to have been introduced into China from the “west”, presumably 

Afghanistan and Iran, about 2100 years ago, which may explain the similarity of types grown in northwestern 

and central areas of China to those grown in Iran and Afghanistan. In both areas spineless, red- or orange-flowered 

types are most common, and in some areas winter types are found—winter types germinate in the fall and go 

through the winter in a rosette stage. 

The northwestern region is referred to as the Xinganning Production Region and the central region as the 

Jiluyu Production Region. In the southeastern region, referred to as the Jiangzhemin Production Region, plants 

are short, early, very spiny, and have dentate leaves and orange flowers. In the southwestern region, referred to as 

the Chuandian Production Region, plants are similar in appearance to those of the southeastern region except for 

the less-dentate leaves. Safflower of the southeastern and southwestern regions of China appears similar in many 

respects to that grown in India. As in India it is grown during the warm, dry winter season. Most likely Indian-type 

safflower spread northward over a long period of time to southern China. In any case, as more is learned about safflower 

in China, revisions will be necessary in safflower centers. 

In most of China safflower is a minor crop being grown as a border row to other crops. In the Xinjiang Autonomous 

Region, where about two-thirds of the safflower is grown, there were large fields, many of them handled by 

machinery except for harvest of the flowers which was done by hand. Harvest of the mature plants was done by 

hand or with a combine. The seeds were processed for oil with modern equipment. 

Because indigenous germplasm in 

China is rapidly being replaced with 

introductions and newly developed 

varieties, and because of the culture of 

safflower over a wide range of different 

environments, it is important that 

an effort be made in the near future to 

collect additional material from that 

country. 

Bullock-driven cart transporting harvested safflower. 9 mi E of Aurangabad, 

Maharashtra State, India, March 1, 1965. (Text and photos from 

Knowles 1965a.) 

9

Preservation of germplasm 

In most cases safflower collections were routed through the diplomatic pouch to the Plant Quarantine 

Section of the USDA. After inspection for diseases and other pests and, if necessary, appropriate treatments, 

the seed was passed to the USDA Germplasm Resources Laboratory in Beltsville, Maryland. There documentation 

of each sample was completed and a Plant Introduction Number (PI number) was assigned. Subsequently safflower 

samples were sent to the USDA Regional Plant Introduction Station at Washington State University in Pullman, 

Washington. There they were kept in cold storage and grown out in a nursery at the first opportunity. Both 

selfed and open-pollinated seed were harvested from each introduction. Before the level of germination decreased 

a second grow-out was made, and again selfed and open-pollinated seed were obtained. This collection is known as 

the World Safflower Collection. 

Breeding programs. An important function of the Regional Plant Introduction Station at Pullman is the maintenance 

of key germplasm of US oilseed breeding programs that have been terminated. To date those include 

programs at UC Davis, the USDA program at Davis, and the University of Arizona program at Tucson. 

Germplasm evaluation 

USDA Regional Plant Introduction Station, Pullman. When safflower introductions are grown out here, always 

with appropriate checks, data are collected on plant development and morphology. Descriptions are standardized 

using descriptors published by the International Board for Plant Genetic Resources, Rome (IBPGR 1983). From 

time to time safflower researchers both in the US and in other countries evaluate the World Collection, usually for 

specific traits. The Regional Plant Introduction Station requests that such information be added to the database 

maintained at Pullman. 1 Periodically, the Introduction Station issues a list of entries in the World Collection which 

includes all evaluation data. 2 

Other stations in the US. The World Collection of Safflower was field evaluated in California for resistance to severe 

infestations of Fusarium wilt caused by Fusarium oxysporum Schlecht. f. sp. carthami Klis. & Hous. (Knowles 

et al. 1968). Resistance was found in 35 introductions from 11 countries. Seed from those introductions were sent 

to all safflower breeders who requested them. Klisiewicz and Urie (1982) identified and registered 14 selections 

from safflower introductions that were resistant to all four races of Fusarium wilt (Table 5). Bockelman (1974) 

found that two genes, a recessive gene at one locus and a dominant gene at a second locus, conferred resistance to 

race 3. 

A second serious disease of safflower was Verticillium wilt caused by Verticillium albo-atrum Reinke & Berth. It 

threatened to eliminate safflower in all areas growing cotton, since safflower was susceptible to the same races of 

the disease that attacked cotton. In 1967 the World Safflower Collection, consisting of approximately 1300 entries, 

was grown in a field known to be infested with wilt (Urie and Knowles 1972). Forty-eight introductions that 

showed some resistant plants came from ten different countries (Table 6). For comparison, the level of infection in 

‘US-10’, a susceptible line used as a check in the screening, was 61%. 

Another serious disease in safflower is Phytophthora root rot, caused by Phytophthora drechsleri Tucker, P. 

parasitica Dast, and P. cryptogea Pethyb. & Laff. It is primarily a disease of irrigated safflower, with the damage 

magnified if the plants have been stressed from lack of water prior to the irrigation or plants stand in water after an 

irrigation. In tests over a 5-year period on heavy clay soil following a crop of rice at the UC Davis Rice Facility, 15 

out of 1547 entries in the World Collection were found to carry some degree of resistance (DaVia et al. 1981). 

1 Editors’ note: Johnson, Stout, and Bradley 1993. 

2 Editors’ note: All data are available on request through the USDA ARS Germplasm Resources Information Network (GRIN), http://www. 

ars-grin.gov/npgs/. 

10

Zimmer and Leininger (1965) tested 1200 introductions and 

selections for rust caused by Puccinia carthami Cda., and found 20 

to have resistance. Introductions with resistance came from India, 

Afghanistan, Iran, Iraq, Turkey, Ethiopia, and Morocco. A high 

level of resistance was found in an accession of C. oxyacantha M. 

Bieb., a wild species, from Iran (Zimmer and Urie 1968). 

USDA at UC Davis. From June, 1976 through 1984, USDA Research 

Agronomist A. Lee Urie was assigned to safflower research 

and based with the USDA oilseeds group at the UC Davis campus. 

In addition to work reported above, several other safflower-related 

objectives were accomplished during that period. 

• The genetic relationships between partial hull safflower 

pericarp type and all known pericarp types was determined 

(Urie 1986). A partial hull safflower plant produces approximately 

40% white seeds and 60% partially dark seeds. 

The dark area of the seeds are due to a reduction in the 

outer layer of the pericarp. This reduction results in a seed 

with low hull content and a high oil content. Partial hull 

is recessive to white (normal) hull and reduced hull but is 

inherited independently of striped hull and thin hull. The 

partial hull gene can be visually identified in filial generations 

thus safflower breeders can easily screen large populations. 

Table 5. Selections of safflower with resistance 

to four races of Fusarium wilt and 

registered with Crop Science Society of 

America.* 

Registration 

number. 

Germplasm 

number 

PI 

number 

Table 6. Summary of number and 

origin of Verticillium wilt-resistant 

introductions.* 

Range of 

Number of resistance 

Country introductions (in %) 

Egypt 5 3–17 

India 9 2–11 

Iraq 1 4 

Iran 15 2–11 

Israel 3 6–8 

Italy 1 4 

Pakistan 1 14 

Philippines 1 2 

Portugal 2 2–6 

Turkey 10 2–3 

*Source: Urie and Knowles 1972. 

Country of 

origin 

18 3992 250010 Iran 

19 4297 250538 Egypt 

20 4305 250608 Egypt 

21 4309 250079 Egypt 

22 4298 250539 Egypt 

23 4343 306596 Egypt 

24 4046 250828 Iran 

25 4011 250827 Iran 

26 4258 253387 Israel 

27 4022 251398 Iran 

28 3133 250830 Iran 

29 3238 209288 India 

30 4043 250823 Iran 

31 3119 250523 Egypt 

*Source: Klisiewicz and Urie 1982 

• Several multiple-disease resistant safflower germplasms 

were developed and released in cooperation with plant pathologists at Davis, California and Beltsville, 

Maryland. A verticillium and fusarium wilt resistant striped hull safflower germplasm was released (Urie 

et al. 1976). A phytophthora root rot resistant germplasm known as 14-5 was released after four years of 

screening in infested field root rot nurseries. This germplasm was one of the 27 families that comprised 

VFstp-l. (Urie et al. 1980). Another lettuce mosaic virus resistant germplasm originating from VFstp-l was 

released (Thomas et al. 1978). 

• When the USDA oilseeds unit was closed out at Davis in 1984, a collection 

of “specialized safflower germplasm” was sent to the USDA 

Regional Plant Introduction Station at Pullman WA. This was a 

diverse collection of about 1800 samples. Included were about 900 

entries from a selected plant introduction nursery, rust-resistant and 

multiple disease-resistant lines, bulks from partial hull progenies, 

VFstp-l families, fusarium-resistant lines from Klisiewicz, pure lines 

with Nebraska and Utah numbers, Mexico Dwarf selections, about 

200 packets of wild species crosses from the Knowles’ program, and 

several other miscellaneous seed lots. 

11

Species relationships in Carthamus 

All wild species of safflower are spiny weeds, some of them very serious because they occupy fields 

sown to other crops. Others are more prevalent in roadsides and waste places. Wild species extend from around 

the Mediterranean Sea to northwestern India. Some species are serious weeds in Australia, and three species, C. 

lanatus L., C. baeticus (Boiss. & Reuter) Nyman, and C. leucocaulos Sibth. & Sm. are minor but potentially serious 

weeds in California. Collections of wild species that were made in the 1958 and 1964–65 missions are listed in 

Appendix 1 Tables 2 and 4, respectively. Species of the genus (wild and cultivated) fall into five groups based on 

chromosome number (10, 11, 12, 22, and 32 pairs of chromosomes). 

Species with 10 pairs of chromosomes. These species are found around the eastern margins of Mediterranean Sea 

and extending eastward into northwestern Iran. Foliage is grey green in color, flowers range from purple to white 

in color, and the pollen is white, but there is a great deal of variation in other morphological characters. Crosses of 

different species indicated that it is a complex taxonomic group (Schank and Knowles 1964). The F1 plants of 

most crosses have normal chromosome pairing, but a translocation differentiates parents of some crosses. 

One species in the group, C. leucocaulos, which was not included in studies of Schank and Knowles (1964), is 

distinctly different from other species. Stems are smooth, head size is small, when colored, flowers are light purple 

or light pink instead of purple, and self-fertility is high. It is endemic to the Aegean Islands, but has been introduced 

to southern Europe, Australia, and the US. Studies of crosses to other species with 10 pairs of chromosomes 

showed a close relationship, though in one cross there was a translocation and paracentric inversion (Estilai 

and Knowles 1978). Crosses to C. nitidus Boiss. with 12 pairs of chromosomes gave a sterile F1 hybrid with very 

irregular chromosome configurations at metaphase I of meiosis. There was very poor chromosome pairing in a 

sterile F1 hybrid of a cross between C. leucocaulos and C. tinctorius. 

Species with 11 pairs of chromosomes. Only one species, C. divaricatus (Beg. & Vacc.) Pamp. is known with this 

chromosome number. It is endemic to Libya and is distinct morphologically. It has horizontal branches; strongly 

divaricate outer involucral bracts; yellow, purple, and white flowers; yellow pollen; and dark-purple-striped 

anthers. It is self-incompatible, and different flower-color types cross readily to give fertile hybrids (Estilai and 

Knowles 1976). It appears that C. divaricatus is closely related to the species with 10 pairs of chromosomes, because 

crosses are easily achieved, the hybrids’ pollen is partially viable, there is good chromosome pairing in PMCs 

of F1 plants, and the F1 hybrids are partially fertile. F1 hybrids with C. tinctorius were vigorous, but produced no 

seed on selfing or in backcrosses to C. tinctorius. C. divaricatus crossed readily with C. lanatus, but the F1 plants 

were sterile. Additional studies are necessary to state positively the origin of C. divaricatus. 

Species with 12 pairs of chromosomes. Cultivated safflower (C. tinctorius) belongs to this group. Three species 

are known to be closely related to cultivated safflower: C. flavescens Spreng, found in continental areas of Turkey, 

Syria, and Lebanon, usually as a weed in wheat fields; C. oxyacantha, found in continental areas from western Iraq 

through northwestern India, and extending northwards to southern USSR, in all areas a very serious weed; and 

C. palaestinus Eig., found in desert areas of Iraq, Jordan, and Israel. C. flavescens is self incompatible (Imrie and 

Knowles 1970), C. palaestinus is self compatible like the cultivated species, and C. oxyacantha is mixed self compatible 

and self incompatible. Two other species, C. gypsicola Iljin, similar to C. oxyacantha and restricted to the 

USSR, and C. curdicus Hanelt, with characteristics of both C. gypsicola and C. flavescens and restricted to northern 

Iraq, obviously belong to the 12-chromosome-pair group (Hanelt 1963). They have not been introduced to the 

US. All of the above species have yellow flowers, except C. palaestinus in which there are both yellow- and whiteflowered 

types. All, including the cultivated species, have yellow pollen. 

Ramanamurthy (1963) found that the self-incompatibility system in C. oxyacantha was of a sporophytic type 

and was determined by multiple alleles at a single locus, termed S, with evidence of dominance between the alleles 

in the pollen. Imrie and Knowles (1971) found a similar situation in C. flavescens, with evidence of considerable 

nonhomology of S alleles in C. flavescens and C. oxyacantha. 

Imrie and Knowles (1970) found in crosses of C. flavescens and C. tinctorius that the following respective differences 

between them were due to alleles at a single locus: long vs. short rosette stage of growth; lobed vs. entire 

12

leaf margins; shattering vs. nonshattering of the seed; pigmented vs. white seed; presence vs. absence of pappus; 

and purple vs. green midveins of the cotyledonary leaves. 

It is believed that all of the above 12-chromosome-pair species originated from a common ancestor, probably in 

the area of northern Iraq and northwestern Iran (Knowles 1976 1 ). All are adapted to an arid or desert climate with 

some winter rainfall. 

A final species with 12 pairs of chromosomes is C. nitidus, which is mainly distributed in the Syrian-Palestine 

region. With white to light rose-colored flowers, white pollen, and grey-green foliage, it is distinctly different from 

other species with 12 pairs of chromosomes. It resembles species of the 10-chromosome-pair group, but in crosses 

with the latter group no seed was produced (Knowles and Schank 1964). However, in crosses with cultivated 

safflower some seeds were produced which gave F1 plants that were intermediate between the parents, and which 

failed to produce seed. Backcrosses to the cultivated species did not produce seed. C. nitidus is self fertile, and its 

taxonomic status is still in doubt. 

Species with 22 pairs of chromosomes. Only one species, C. lanatus, has 22 pairs of chromosomes. It is mostly 

yellow flowered, but there are some white-flowered types. It has yellow pollen and is self fertile. Its range is very 

wide, extending in the Old World from Spain 

and Portugal on the west, around the Mediterranean 

Sea and eastward into eastern Turkey. 

At first it was assumed that it was an alloploid 

resulting from natural crosses of species with 10 

and 12 pairs of chromosomes. However, a study 

by Harvey and Knowles (1965) included 

crosses of species of the two groups and artificial 

alloploids developed from their F1 hybrids, 

crosses of artificial alloploids to C. lanatus and 

diploid species, and crosses of 10- and 12-chromosome-pair 

species to C. lanatus. Morphological 

characters and chromosome behavior at MI 

did not provide convincing evidence that any of 

the 10- and 12-chromosome-pair species of the 

study were ancestral species of C. lanatus. Also, 

A 

in an earlier study (Ashri and Knowles 1960) 

with only four species with 10 pairs of chromosomes, 

and three with 12 pairs, inconclusive 

results were obtained. Further study is necessary 

to ascertain the origin of C. lanatus. 

Species with 32 pairs of chromosomes. There 

are two species in this group, C. turkestanicus M. 

Popov and C. baeticus. C. turkestanicus ranges 

from Turkey on the west to northern Pakistan 

and Kashmir on the east, with presumably introduced 

populations in Ethiopia. C. baeticus may 

be found around the entire Mediterranean Sea 

and on its islands. As an introduced weed, it is 

found in California 2 and British Columbia. 

Both species have grey-green foliage, light 

yellow to white flowers, light-colored anthers 

often with purple to brown stripes, and white 

pollen (Khidir and Knowles 1970a). C. 

1 Editors’ note: Updated in: Knowles and Ashri 1995. 

2 Editors’ note: As C. creticus L. 

13 

B 

Threshing safflower (A, south of Buldana, Maharashtra State, 

India, February 24, 1965) and winnowing safflower (B, near 

Bhir, Maharashtra State, India, March 4, 1965). (Text and photos 

from Knowles 1969a).

aeticus has less pubescence; fewer, longer, and narrower involucral bracts; narrower heads with fewer florets and 

achenes; and shorter corolla lobes. C. turkestanicus resembles C. lanatus more closely than does C. baeticus, but 

compared to C. lanatus it has wider bracts, a more distinct floret saccation, less pubescence, and white instead of 

yellow pollen. Chromosome pairing of both C. turkestanicus and C. baeticus was regular, but hybrids between them 

had a few trivalents and quadrivalents. 

Alloploids synthesized from C. leucocaulos × C. lanatus were similar to C. baeticus (Khidir and Knowles 

1970b). Because the parental species overlap in the eastern Mediterranean, it is assumed that C. baeticus had its origin 

there. The situation was similar for alloploids developed from crosses of C. glaucus ssp. glaucus (I.B.) Schrank 

by C. lanatus; they were similar to C. turkestanicus. It is assumed that C. turkestanicus originated in Transcaucasia, 

just west of the Caspian Sea where the two species overlap (Hanelt 1963). 

Both C. baeticus and C. turkestanicus cross readily with C. lanatus, the hybrids having about 40% stainable 

pollen and producing some selfed seed. Similar results were obtained by Ashri and Knowles (1960). Where C. 

lanatus and C. baeticus occur together on the island of Crete, the two species are similar in appearance except for 

pollen color (Khidir and Knowles 1970b). The same is true of C. lanatus and C. turkestanicus in Thrace. This suggests 

that there is considerable gene exchange between C. lanatus and the species with 32 pairs of chromosomes. 

Evolution in the genus. A tentative pattern of evolution in the 

genus Carthamus is presented in Figure 2. Khidir and Knowles 

(1970b) suggested the genomic formula A2A2 for C. leucocaulos, 

but later studies (Estilai and Knowles 1978) indicated that the 

chromosomal differences between A2A2 and AA did not warrant 

different formulas. A genomic formula was not assigned to C. nitidus 

with 2n=24 nor to C. divaricatus with 2n=22. However, the evidence 

from the studies of Estilai and Knowles (1976) would indicate 

that it resulted from a cross between a species with 2n=20 and either 

the species C. lanatus (A1A1B1B1) or a species with the genomic 

formula BB (2n=24). 

Wild species in the improvement of C. tinctorius. With the exception 

of C. nitidus, C. tinctorius has crossed readily with wild species 

having 12 pairs of chromosomes, to give fertile F1 and F2 progeny. 

The fact that F1 plants have been obtained from crosses of C. tincto- 

rius to species with 10 pairs of chromosomes (Ashri and Knowles 1960; Estilai 1977; Estilai and Knowles 

1978), with C. divaricatus with 11 pairs of chromosomes (Estilai and Knowles 1976), with C. lanatus with 22 

pairs of chromosomes (Ashri and Knowles 1960; Heaton and Klisiewicz 1981; Hill 1981), and with C. nitidus, 

a distantly related species with 12 pairs of chromosomes (Knowles and Schank 1964) indicates that these 

distantly related species may prove to be a source of useful genes. 

From a cross of a wild species as the female parent to C. tinctorius (A.B. Hill, personal comm.) cytoplasmically 

male-sterile plants were obtained. Heaton and Klisiewicz (1981) produced an alloploid from a cross of C. tinctorius 

and C. lanatus. Tested against diseases to which C. tinctorius is susceptible, bacterial blight caused by Pseudomonas 

syringae van Hall, leaf spot caused by Alternaria carthami Chowd., and wilts caused by Fusarium oxysporum f. 

sp. carthami and Verticillium albo-atrum, C. lanatus and the alloploid were resistant or highly resistant. The authors 

speculated on the use of the alloploid in breeding programs. 

Among a number of wild species that were examined for the fatty acid composition of the seed oil, none was 

found to be markedly different from high linoleic C. tinctorius (Knowles 1972). 

14 

A 1 A 1 

B 1 B 1 

AA A A B B BB 

1 1 1 1 

AAA A B B 1 1 1 1 

Figure 2. Genome evolution in genus 

Carthamus. AA is the assumed genome formula 

for species with 2n=20; A1A1 for an 

ancestral species with 2n=20; BB for species 

with 2n=24; and B1B1 for an ancestral 

species with 2n=24.

Fatty acid composition of safflower oil 

For several years safflower germplasm was examined for fatty acid using the iodine value which was 

measured using a hand refractometer. The iodine value gave a reasonably accurate measure of the saturation of the 

oil (Knowles 1965b), as there was some variation in two fatty acids, oleic and linoleic. Iodine values varied between 

138 and 145, with fatty acid compositions averaging about 76% linoleic, 16% oleic, 1% stearic, and 7% palmitic 

(Knowles and Mutwakil 1963). 

Several introductions from India were found in 1957 to have iodine values much lower than that of the standard 

type. It turned out that these were mixtures of the standard type and a mutant type which had iodine values 

averaging about 90, a complete reversal in the amounts of oleic and linoleic acid, and little change in the amounts 

of stearic and palmitic acids. It was found (Knowles and Mutwakil 1963) that the difference in levels of linoleic 

and oleic acids was governed by one gene. 

Through backcrossing, the high oleic version of the gene was transferred from the mutant type to US-10, to give 

a commercially acceptable high oleic cultivar that was named UC-1 (Knowles et al. 1965). Though the oilseed 

industry showed little interest in the mutant type when first discovered, its attitude had changed when UC-1 was 

developed. UC-1 was grown commercially for a few years, but was soon replaced by cultivars developed by seed 

companies that had used UC-1 in their breeding programs. 

The discovery of the high oleic gene, termed ol, meant that there were two totally different oils available from 

safflower. The high linoleic type was used in many food preparations because of its high polyunsaturation. It was 

also a useful oil in the manufacture of varnishes, paints, and similar products. On the other hand, the high oleic 

type, being monounsaturated and heat stable, has had increasing use in the food industry. Its chemical composition 

is similar to that of olive oil, but it has a bland taste. Details of this development are recorded in “The Errant 

Gene”, a chapter in the book Garden in the West (Wells 1969). 

Subsequently, another mutant was found, this one with about equal amounts of oleic and linoleic acids, and 

with no change in levels of palmitic and stearic acids. It was found in the town of Mianeh in Iran—interestingly, 

“mianeh” is the Iranian word for intermediate—and was assigned PI 254717 by the USDA. A study of its inheritance 

(Knowles and Hill 1964) showed that it was caused by the allele ol 1 , at the same chromosome locus as ol. 

The oilseed industry had little interest in this intermediate type of oil, since it could be constituted by a blend of 

equal amounts of the high linoleic and high oleic oils. Furthermore, the seed oil proved to be sensitive to temperature: 

high temperatures during the seed development period giving an oil high in oleic acid, and cool temperatures 

giving a high linoleic oil (Bartholomew 1971). 

Third and fourth mutants were found in introductions PI 226993 from Israel and PI 262430 from Russia, 

respectively. They each had seed oil with stearic acid levels of 5 to 12% instead of the usual levels of 1.0 to 2.5%. An 

inheritance study (Ladd and Knowles 1970) showed that genes at one locus governed the difference—both mutants 

had the genotype stst and the standard low stearic acid type had the genotype StSt. The st locus was independent 

of the ol locus (Ladd and Knowles 1971). The high stearic type was of no commercial interest because levels 

were low, and stearic acid could be obtained more cheaply from other sources. 

A study by Hill and Knowles (1968) showed that the genes governing fatty acid composition showed their 

effects about 10 days after flowering, when oil content and seed weight were increasing at a rapid rate. 

A final fatty acid mutant was found in an introduction from Portugal (PI 253568). This introduction had 

linoleic acid contents of the seed oil between 85 and 90%, some 10% higher than the standard high linoleic type 

grown commercially. The levels of all other fatty acids were reduced. One gene, termed li, at a different locus from 

the ol and st genes, was responsible for the very high levels of linoleic acid (Futehally 1982). 

The UC Davis experience with fatty acid composition of safflower clearly shows the value of making collections 

of safflower from many geographic areas. In some cases, because of the very small populations providing mutants, 

germplasm collection was a rescue operation that preserved genes that may have future commercial value. 

15

Male sterility 

A very thin-hulled mutant of safflower was discovered by Rubis (1962) which was governed by the recessive 

gene th. The mutant trait was associated with weak stems and much-delayed pollen release. It was believed 

that delayed pollen release would serve as a form of structural male sterility. Efforts by plant breeders to develop 

genotypes to develop usable structurally male-sterile genotypes were unsuccessful. The delay in, and amount of, 

pollen release was unpredictable, so there could be an unpredictable amount of thin-hulled types, which were low 

yielding in hybrid varieties. Efforts were made through selection to separate the thin-hulled trait from weak stems, 

delayed pollen release, and low yield, but without success. 

Ebert and Knowles (1968) found that the genotype thth for thin hulls had one primary effect: it reduced or 

prevented secondary wall formation in the cells of “fibrous” tissues. This resulted in thin hulls of the seeds, weak 

stems and branches, and delayed dehiscence of the anthers—th was a gene with a pleiotropic effect. The absence 

of secondary wall formation in the anthers, which is involved in dehiscence, resulted in a collapse of the walls in a 

disorganized manner, resulting in failure or delay in the release of pollen. 

Heaton and Knowles (1982) reported the discovery of genetic male sterility in accession PI 263914 from 

Afghanistan that had been treated with colchicine. It was governed by a single recessive gene ms. Two germplasm 

releases were made, UC-148 with orange flowers, developed from the original mutant, and UC-149 with white 

flowers developed from a cross of UC-148 with PI 340088, an introduction from Turkey (Heaton and Knowles 

1980). 

Genetic male sterility was not acceptable for commercial use in the US because of labor costs necessary to remove 

fertile (Msms) genotypes from the female parent (a 1:1 mixture of Msms and msms). In India, however, it is 

being used by one company which is experimenting with a female parent that has appressed branches (apap) and 

is spineless—this makes removal of the male-fertile types much easier. If the male parent has spreading branches 

and spines, the F1 hybrid will be the same. 

A 

16 

B 

The first step in processing safflower for oil in Egypt was 

cracking seed with a stone grinder sufficiently to loosen the 

hulls. Screens were used to separate hulls from seed. The 

second step was grinding of the seed with a rolling round 

stone, pictured in A (Large roller used to crush oil seeds. 

Assiut, Upper Egypt. June 19, 1958). Finally this ground seed 

was placed in tight flat straw baskets shaped like a beret pictured 

in B (Flat basket made of reeds. Assiut, Upper Egypt. 

June 19, 1958). Several filled baskets were stacked one on top 

of the other and pressure applied to the top with either a 

hand-operated screw press or a heavy wooden lever. About 

125 kilos of seed gave 20 kilos of oil. (Text and photos from 

Knowles 1959.)

Already mentioned are the cytoplasmic male-sterile genotypes developed by AB Hill. They are not yet being 

used commercially. 

In many crosses of distantly related safflower species, sterile types appear in the F2. Carapetian and 

Knowles (1976) studied such sterile types and found that the sterility was governed by three genes, tentatively 

identified as a, b, and c. Sterile genotypes were aabbC_, aaB_cc, and aabbcc; the expected ratio in F2 was 57 fertile 

to 7 sterile. A study by Carapetian indicates that a is linked to ol, the gene causing high levels of oleic acid in the 

seed oil. 1 

1 Editors’ note: Now published: Carapetian and Knowles 1993. 

Miscellaneous traits 

In the course of germplasm evaluation and inheritance studies over several years, several mutants 

have been identifed. Some were studied in detail, others not, and they will be briefly discussed. Some studies 

included introductions different from commercial cultivars. 

Branching attitude. Several introductions, most of them from India, had branches appressed against the main 

stem, the angle of branching was not above 20°. Commercial cultivars had branch angles above 40° and rarely 

above 80°. One recessive gene, ap, governed the appressed trait (Leon and Knowles 1964). 

A mutant was found with decumbent branches, where the angle of branching was 80 to 90° and sometimes 

exceeded 90°. Temple and Knowles (1975) found that one recessive gene, dec, was involved. Fernández and 

Knowles (1978) found that dec was independent of ap. The genotype ap/ap, dec/dec had spreading branches 

indicating the genes counteracted one another. 

Closed flowers. Three different closed-flower types were identified and were termed arrowhead (Type A), bar 

(Type B), and cage (Type C). The petal lobes of Type A are closed in the upper half and separated slightly in the 

lower half. The style projects through the fused portion and attains normal length. Seed set is normal. In Type B 

the petal lobes are strongly fused through their entire length, and the style and anther tube do not emerge. Seed set 

is greatly reduced. In Type C only the tips of the corolla lobes are fused which prevents emergence of the style and 

anther tube, though the latter may emerge laterally. Seed set is reduced about 50%. Dillé and Knowles (1975) 

found that fusion of petal lobes was due to epidermal cells of the margins of lobes being differentiated into papillae 

with pointed appendages that extend into or mesh with papillae of adjacent lobes. Limited data from inheritance 

studies indicated that independent single recessive genes were responsible for the development of the aberrant 

papillae. 

Brittle stems. A brittle-stemmed safflower mutant appeared in the F6 of a cross between an introduction from Turkey 

(PI 175624) and ‘Pacific 7’. It was weak and droopy in appearance. When bent slightly the branches and main 

stem broke cleanly, so that at maturity there were many branches lying on the ground. Hulls were frequently soft 

and bubbly in appearance. A single recessive gene br was involved (Temple and Knowles 1975). Limited chemical 

analysis showed that brittle plants had lower crude fiber, higher protein content, and higher levels of starch and 

soluble polysaccharides than did tough-stemmed cultivars. 

Black foliage. The black mutant, which appeared in the breeding nursery, expressed the trait as the plants approached 

maturity. Only leaves and bracts were involved. When homozygous, one recessive gene, bl, resulted in a 

black plant (Temple and Knowles 1975). 

Crinkle leaves. A crinkle-leaved mutant was found to be due to a single recessive gene, cr (Temple and Knowles 

1975). 

Light-green leaves. The light-green mutant was due to a single recessive gene that was not assigned a symbol 

(Temple and Knowles 1975). 

17

Flower color. Hartman 

(1967) identified a corollacolor 

type in safflower based 

on emergent flower color 

just as the flower emerged 

from the bud and before the 

lobes open, and mature color 

obtained as flowers wilted 

when flower color was most 

intense. Fresh flower color was 

obtained when the flowers 

had opened but not yet wilted. 

Descending paper chromatography 

was used to separate 

pigments in fresh flowers that 

had been dried immediately 

and the petals ground to a 

powder. The flower colors 

were identified under natural 

and long-wave ultraviolet 

(UV) radiation. Three main 

pigments were identified 

under natural light: carthamin 

Table 7. Association of flower colors of safflower with chromatography colors. 

Emergent color 2 Mature color 2 Carthamin 2 

Chromatography colors 

Under natural light 

Safflower 

yellow 

Yellow No. of spots under 

pigment3 UV radiation 

white white 0 0 3 

cream near white 5 4 0 3 5 

sulfur sulfur 0 4 3 0 

pc yellow pink 3 2 3 0 

pc yellow light orange 2 4 3 2 

c yellow yellow 0 10 3 0 

l yellow orange 4 6 3 0 

l yellow orange red 5 

orange red 10 10 3 0 

1 On a scale of 0 (none) to 10 (intense). 

2 Flower and chromatography colors: c = canary; l = lemon; pc = pale canary. 

3 Unidentified. 

4 Straw yellow, not red. 

5 Same as red but amounts of carthamin reduced by an unspecified amount. 

which remained as a reddish spot at the site of the placement of the pigment extract and safflower yellow and 

an unidentified yellow pigment, both of which separated out near the advancing edge of the solvent. Under UV 

radiation several additional colors were identified. Table 7 associates flower colors with colors identified under UV 

radiation. The chemistry of carthamin and safflower yellow are well known. Before the development of aniline dyes 

in the last century, carthamin, from safflower, and indigo were the most important sources of dye for textiles. The 

uses in China today of carthamin, safflower yellow, and dried petals have already been mentioned. 

On the basis of F2 analyses of crosses of the different flower color types, Hartman (1967) identified four loci 

that governed flower color: c with three alleles, y with three, o with two, and r with three. 

Thin hull. There has been a gradual increase in oil content in commercial cultivars of safflower from about 37% in 

1950 to about 43% in the 1970s. This was achieved mostly by reduction in the amount of seed hull. A very thinhulled 

mutant has already been mentioned as a possible female parent for hybrid cultivars—it was not acceptable. 

Urie and Zimmer (1970) found types which they designated reduced hull that increased oil content from 34% 

for US-10 to an average of 42.7%. Later a partial-hull mutant was found (Urie 1986) that reduced hull content still 

further—the six original mutant plants had an oil content of 49.0%. In yield trials the yield of partial hull and the 

average yield of two commercial cultivars were 1688 and 1668 lb/ac (1890 and 1868 kg/ha), respectively. One recessive 

gene, par, governed the expression of partial hull. It should prove to be a useful gene in breeding programs. 

18

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USDA Circ No 366. Government Printing Office, Washington DC USA. 

rAmAnAmUrthy GV 1963. Relationships of cultivated safflower (Carthamus tinctorius L.) to the wild species C. oxyacantha 

M.B., PhD Dissertation, Univ California, Davis CA USA. 

rUBis DD 1962. A mutant gene for thin hull in safflower. page 75 in: Agron Abs, 54th Annual Mtg, Am Soc Agron. 

Madison WI USA. 

schAnK SC and PF Knowles 1964. Cytogenetics of hybrids of Carthamus species (Compositae) with 10 pairs of chromosomes. 

Am J Bot 10:1903–1102. 

shinn CH 1903. Culture Work at the Substations 1899–1901. Univ California Agric Exp Sta Bull 147. Berkeley CA 

USA. 

temple SR and PF Knowles 1975. Inheritance of brittle stems in safflower. Crop Sci 15:694–697. 

thomAs CA, LH zimmermAn, and AL Urie 1978. Registration of LMVFP-1 safflower germplasm. Crop Sci 18:1099. 

Urie AL 1986. Inheritance of partial hull in safflower. Crop Sci 26:493–498. 

Urie AL, DJ DAViA, PF Knowles, and LH zimmermAn 1980. Registration of 14-5 safflower germplasm (Reg. No. GP 15). 

Crop Sci 20:115–116. 

Urie AL and PF Knowles 1972. Safflower introductions resistant to Verticillium wilt. Crop Sci 12:545–546. 

Urie AL and DE zimmer 1970. A reduced-hull seed character of safflower. Crop Sci 10:371–372. 

Urie AL, LH zimmermAn, CA thomAs, WF peterson, and PF Knowles 1976. Registration of VFstp-1 safflower germplasm 

(Reg. No. GP 12). Crop Sci 16:888. 

VAViloV NI 1951. The origin, variation, immunity and breeding of cultivated plants. [Transl. K.S. Chester] in: Selected 

Writings of N.I. Vavilov. Chronica Bot 13(1/6). 364 pages. 

wells GS 1969. Garden in the West. Dodd, Mead, & Co, New York NY USA. 

wU Y-S and D li 1981. Distribution, utilization, and history of cultivation of safflower in China. pages 272–275 in: 

PF Knowles (ed) Proceedings of the First International Safflower Conference, Davis, California, July 12–16, 1981. Univ 

California, Davis CA USA. 

zimmer DE and LN leininger 1965. Sources of rust resistance in safflower. Pl Dis Reptr 49:440–442. 

zimmer DE and AL Urie 1968. Inheritance of rust resistance in crosses between cultivated safflower, Carthamus tinctorius 

L., and wild safflower, Carthamus oxyacantha M.B. Phytopath 58:1340–1342. 

21

Appendix 1 

Appendix 1. Table 1. Count of accessions of cultivated safflower 

(Carthamus tinctorius) collected in 1958*. 

Country Area Count Remarks 

India Pusa Station 30 

Hyderabad 88 34 m from Rajenderonayer Sta. 

Poona 0 Seed samples promised later 

Amritsar 7 From bazaars 

Delhi 0 

Total 125 

Pakistan Lahore 9 All from bazaars 

Lyallpur 14 From Lyallpur Station 

Gujerat 24 From bazaars and farmers 

Rawalpindi 13 From bazaars 

Peshawar 3 From bazaars 

Karachi 12 From bazaars 

Total 75 

Afghanistan Kandahar 0 

Lashkar Gah 0 

Kabul 15 From bazaars and fields 

Ghazni 1 From a border of a field 

Total 16 

Iran Tehran 10 From bazaars and Karaj Sta. 

Khuzestan 0 

Isfahan 22 From bazaars and fields 

Tabriz 7 From bazaars and homes 

Mashad 14 From bazaars and fields 

Total 53 

23

Appendix 1. Table 1. Continued.* 

Country Area Count Remarks 

Iraq Baghdad 4 From bazaars 

Mosul 0 

Sulaimaniya 0 

Total 4 

Jordan Amman Jerusalem 13 Mostly from fields 

Syria Damascus Aleppo 6 From bazaars 

Israel Near Jerusalem 1 From a villager 

Near Nazareth 3 From villagers 

Total 4 

Turkey Univ. of Ankara 3 

Eskişehir Exp.Sta. 7 

Eskişehir area 4 Farmers’ fields 

Belikesir area 1 Farmers’ fields 

Total 15 

Greece Larissa Station 11 From Turkey 

Egypt Cairo 7 From bazaars 

Giza Station 2 

Near Giza 2 

Alexandria 2 Bazaar and Exp. Sta. 

Kena 15 Warehouses 

Near Luxor 1 Farmer 

Sohag 4 Oilseed mill and farms 

Assiut 3 Bazaar and oilseed mill 

Total 36 

Morocco 0 

Spain Cuenca area 5 Jefatura Agronomica 

Teruel area 2 Villagers 

Alicante 1 Seed store 

Elche 2 Seed store 

Southern Spain 0 

Total 10 

Portugal Southern part 10 Sacavem Station 

Herbaria of Asia 

and Europe 

52 Sacavem Station 

Total 62 

Total all countries 

*Source: Knowles 1959. 

420 

24

Appendix 1. Table 2. Count of accessions of wild species of safflower 

(genus Carthamus) collected in 1958*. 

Species/country Count Species/country Count 

C. flavescens C. alexandrinus 

Iraq 2 Egypt 4 

Syria 7 C. lanatus 

Turkey 8 Afghanistan 5 

C. oxyacantha Egypt 2 

Afghanistan 12 Iran 14 

India 14 Iraq 5 

Iran 44 Morocco 15 

Iraq 42 Spain 15 

Pakistan 68 Turkey 15 

C. nitidus Pakistan 9 

Jordan 2 Portugal 3 

Syria 2 C. baeticus 

C. dentatus Morocco 4 

Turkey 17 Spain 2 

C. glaucus C. palaestinus 

Iran 9 Greece 17 

Iraq 13 Iraq 4 

Israel 4 C. arborescens 

Jordan 3 Spain 2 

Syria 1 C. caeruleus 

Turkey 7 Morocco 5 

C. syriacus Turkey 1 

Jordan 9 Species hybrids 

Syria 2 Iran 3 

C. tenius Iraq 3 

Israel 15 Israel 11 

Jordan 2 Jordan 4 

Syria 3 

Turkey 2 Total 431 

*Source: Knowles 1959. 

25

Appendix 1. Table 3. Count of accessions of cultivated safflower (Carthamus tinctorius) 

collected in 1964–65*. 

Country Area or location Date 

Type of 

collection Count Use 

Turkey Eskişehir 7/29-30 Seed 3 Oil 

Eskişehir Res. Sta. 7/30 Seed 12 Oil 

Eskişehir to Ankara 7/30 Heads 2 Oil 

Ankara Univ. 7/31 Seed 2 Oil 

İskenderun–Antakya– 

Reyhanlı 

8/5 Seed & heads 2 Flowers 

Gaziantep–Maras 8/6-7 Seed & heads 8 Flowers 

Total 29 

Iran Azerbaijan 9/2-6 Seed & heads 15 Flowers & seed 

Teheran–Isfahan–Shiraz 9/7-17 Seed 25 Flowers & seed 

Gorgan–Mashad 9/19-21 Seed 2 Flowers & seed 

Total 42 

Afghanistan Herat bazaar 9/24 Seed 6 Flowers & seed 

Kabul–Ghazni 9/26-29 Heads 3 Oil 

Total 9 

Pakistan Lyallpur bazaar 10/5 Seed 1 Seed 

Bangladesh Dacca area 11/23 Seed 5 Seed 

India Kashmir 10/10-11 Seed 13 Flowers & seed 

Pathankot area 10/13 Seed 3 Seed 

Nainital area 10/28 Seed 1 Seed 

Allahabad Agric. Inst. 11/8 Seed 2 Oil 

Banares Hindu Univ. 11/12 Seed 1 Oil 

Muzaffarpur, Bihar State 11/15 Seed 40 Oil & vegetable 

Sabour area, Bihar State 11/18 Seed 11 Vegetable 

Mysore State 1/5-2/9 Seed & heads 25 Oil 

Andhra Pradesh State 1/21-/2/8 Seed & heads 75 Oil 

Maharashtra State 2/9-3/4 Seed & heads 218 Oil & vegetable 

Total 389 

Sudan Univ. of Khartoum and 

Hudeiba Res. Sta. 

3/30-4/1 Seed & heads 18 Roasted seed 

Egypt Upper Egypt 4/14-22 Seed & heads 23 Oil & flowers & vegetable 

Spain Cordoba area 5/30 Seed & heads 1 Oil 

Elche area 6/12 Seed & heads 2 Flowers 

Total 3 

Total all countries 

*Source: Knowles 1965a. 

519 

26

Appendix 1. Table 4. Accessions of wild species of safflower (genus Carthamus) collected in 1964–65*. 

Species/country and area Collection number Species/country and area Collection number 

C. flavescens C. lanatus 

Turkey, eastern 146, 149, 157, 164, 212, 214, 215, 216 Algeria 898, 910, 911 

Turkey, southern 103, 104, 106–111, 127, 128, 129, 

131, 135, 138 

Crete 17, 18, 27 

Turkey, western 94, 96, 97, 99 Greece, northern 46, 48, 51, 53, 54, 58, 59, 63 

C. oxyacantha Greece, southern 40, 41 

Iran, northeast 275, 276 Libya 871, 872, 875, 876, 877 

Iran, southern 264, 265 Morocco 922, 926 

Iran, western 218, 222, 232, 237, 238, 239 Turkey, Thrace 66 

C. dentatus Turkey, western 70, 73 

Crete 6, 9 Tunisia 878, 885, 892 

Greece, northern 50, 55, 57, 64 Yugoslavia, Adriatic 1, 2, 3 

Greece, southern 38, 39, 42 Yugoslavia, southern 4 

Rhodes 32, 33, 34, 35 C. lanatus × C. dentatus 

Turkey, eastern 163, 213 Greece, northern 56 

Turkey, southern 102, 112, 123, 126 C. turkestanicus 

Turkey, Thrace 65 Iran, northeastern 274 

Turkey, western 69, 75 Iran, western 217, 224, 240 

Yugoslavia, southern 4 Turkey, eastern 203, 211 

C. dentatus × C. glaucus Turkey, Thrace 67, 68 

Turkey, southern 125 Turkey, western 71 

C. glaucus × C. flavescens C. baeticus 

Turkey, eastern 150, 161 Greece, northern 47, 49 

Turkey, southern 134 Morocco 929 

C. tenuis Portugal 940 

Rhodes 297, 307, 317, 377 Turkey, southern 114 

Turkey, southern 1137 C. creticus 

C. alexandrinus Crete 8, 10, 11, 12, 14, 15, 16, 20, 26, 28 

Egypt 863, 875 Rhodes 36 

Libya 868, 873 C. arborescens 

C. leucocaulos Spain 933 

Crete 13, 25 Unknown species 

Greece, northern 52 Libya 869 

*Source: Knowles 1965a 

27

Appendix 1. Table 5. Accessions of Carduncellus species collected in 

1964–65*. 

Species/country 

C. caeruleus (syn. Carthamus caeruleus) 

Collection number 

Algeria 895, 899, 902–907, 909, 912, 913, 917 

Crete 8 

Morocco 923, 928, 931 

Portugal 941 

Spain 932, 934, 935, 936, 937, 939 

Tunisia 

C. calvus 

891 

Morocco 

C. ariocephalus 

920, 921 

Egypt 866 

Tunisia 

C. helenoides (syn. Carthamus helenoides) 

879 

Algeria 

C. hispanicus 

916, 918, 919 

Spain 

C. ilicifolius 

958, 963 

Algeria 

C. mareoticus 

901 

Egypt 861 

Libya 

*Source: Knowles 1965a. 

867 

28

Appendix 1. Table 6. Safflower germplasm selected in China in 1988*. 

No. PI no. † Source 

Seed supplied by Dr. Li Dajue from his safflower nursery at the Beijing Botanical Garden, Institute of Botany. 

11 544015 Kunming City, Yunnan Province 

14 657784 Tacheng County, Xinjiang Province 

19 544016 Zhangye County, Gansu Province 

26 576979 Xiapu County, Fujian Province 

27 544017 Ruicheng County, Shanxi Province 

45 544018 Xiapu County, Fujian Province 

54 544019 Yanjin County, Henan Province 

82 653135 Jinxian County, Hebei Province 

89 544020 C. lanatus from Holland 

90 544021 Botanischer Garden, University of Frankfurt, West Germany 

94 576980 Milan, Italy 

96 544022 Botanical Garden, Liege University, Belgium 

141 544023 Fukang County, Xinjiang Province 

142 543974 Hetian County, Xinjiang Province 

151 576984 Yinchuan City, Ningxia Province 

157 576985 Jardins Botaniques de la Ville et de l'Université Nancy, Meurthe-et-Moselle, France 

161 576986 Hongze County, Jiangsu Province 

163 576987 Mengzi County, Nunnan Province 

168 576988 Huaxian County, Henan Province 

177 576983 Qinghe County, Hebei Province 

180 543975 Nanxi County, Sichuan Province 

182 543976 Heze County, Shandong Province 

183 543977 Zhecheng County, Henan Province 

187 653133 Wuhu City, Anhui Province 

194 543978 Yingshan County, Sichuan Province 

200 543979 Tongliao City, Inner Mongolia 

201 543980 Qiubei County, Yunnan Province 

203 543981 Botanical garden, Johannes Butenberg University, Mainz, West Germany 

220 543982 Kalaqinzuoyi County, Liaoning Province 

224 543983 Zhangye County, Gansu Province 

225 543984 Qixian County, Henan Province 

226 543985 Huolu County, Hebei Province 

227 543986 Danxian County, Shandong Province 

228 543987 Dangshan County, Shandong Province 

229 543988 Qixian County, Henan Province 

232 543989 Gaotang County, Shandong Province 

233 576982 Yongnian County, Hebei Province 

235 543990 Fengqui County, Henan Province 

* Source: Knowles 1989. 

† PI designations are from subsequent accessioning in 1989 by USDA Regional Plant Introduction Station, Pullman WA USA. 

29



237 543991 Boxian County, Anhui Province 

238 543992 Juye County, Shandong Province 

239 543993 Lingquan County, Anhui Province 

240 543994 Caoxian County, Shandong Province 

242 543995 Jingxian County, Hebei Province 

243 543996 Weishan County, Shandong Province 

244 543997 Daming County, Hebei Province 

245 543998 Liangshan County, Shandong Province 

249 543999 Sixian County, Anhui Province 

252 653134 Chengwu County, Shandong Province 

296 544000 Ningling County, Henan Province 

297 544001 Taikang County, Henan Province 

299 544002 Yucheng County, Henan Province 

301 544003 Linzhang County, Hebei Province 

306 544004 Czechoslovakia 

317 544005 Xiuxian County, Anhui Province 

321 544008 Lingxian County, Shandong Province 

323 544009 Donghai County, Jiangsu Province 

324 544010 Changyi County, Shandong Province 

326 544011 Ningjin County, Shandong Province 

327 544012 Guangde County, Anhui Province 

331 544013 Xinchang County, Zhejiang Province 

332 544014 Shanghe County, Shandong Province 

337 544024 Shucheng County, Anhui Province 

338 544025 Huaiyuan County, Anhui Province 

341 544026 Xianxian County, Hebei Province 

342 544027 Xingtang County, Hebei Province 

344 544028 Yuhuan County, Zhejiang Province 

346 544029 Liuan County, Anhui Province 

348 544030 Chengxian County, Zhejiang Province 

350 544031 Anji County, Zhejiang Province 

351 544032 Zouping County, Shandong Province 

356 544033 Kuerle City, Xinjiang Autonomous Region 

369 544034 Fukang County H1-001, Xinjiang Autonomous Region 

375 544035 Fukang County H1-002, Xinjiang Autonomous Region 

383 544036 Yutian County, Xinjiang Autonomous Region 

400 544037 Jimushaer County, Xinjiang Autonomous Region 

407 544038 Tongjiang County, Sichuan Province 

416 544039 Changning County, Yunnan Province 

417 544040 Dahongpao, Yanjin County, Henan Province 



30



418 544041 Xizang, Lasa City, Tibet 

424 544042 Ledu County, Qinghai Province 

444 544043 Taixing County, Jiangsu Province 

445 544044 Nanjing City, Jiangsu Province 

479 544045 Funing County, Jiangsu Province 

480 544046 Lingquan County, Anhui Province 

495 544047 Jiangyin County, Jiangsu Province 

496 544048 Wuwei County, Anhui Province 

509 544049 Taiping County, Anhui Province 

511 544050 Wuxi City, Jiangsu Province 

593 544051 Chuanhong-1, Chongqing City, Sichuan Province 

629 576989 Weishan County, Yunnan Province 

2155 544052 ZW-971, Zhangye County, Gansu Province 



Seed from other sources visited by PF Knowles. 

C-1 544054 Hangzhou Pharmaceutical Institute 

C-2 544063 Qinghai Academy of Agriculture and Forestry, Xining 

C-3 544062 Qinghai Academy of Agriculture and Forestry, Xining 

C-4 544055 From Prof. Wang Zhao-mu, Institute of Industrial Crops, Urumqi, Xinjiang Autonomous 

Region, 7/28/88. Local type XJ-12 











C-10 544061 Seed used for planting on the Red Flag farm, 30km S of Jimusaer, NE of Urumqi, Xinjiang 

Autonomous Region. Spineless type with red or orange flowers, used as a source of 

flowers to obtain both yellow and red pigments. Seed from 1987 crop obtained 7/23/88. 



31

Appendix 2. 

Publications of P.F. Knowles on oilseed crops. 

1949 Houston, B.R. and P.F. Knowles. Fifty-year survival of flax fusarium wilt in the absence of flax culture. Pl Dis 

Reptr 33(1):38. 

1949 Knowles, P.F. Castor beans studied for potential values as oil crop for California production. Calif Agric 

3(3):13–14. 

1949 Knowles, P.F. Dual purpose safflower seed. Produces drying oil for the paint industry and seed meal for 

livestock and poultry feed. Calif Agric 3(2):11,16. 

1949 Knowles, P.F. Safflower, a new crop. Univ. of California Agric Ext Service leaflet. Berkeley CA USA. [20 

pages] 

1949 Knowles, P.F. Safflower production in California. Univ. of California Agr. Expt. Sta. Ext. Service Leaflet. 

Berkeley CA USA. [7 pages] 

1949 Knowles, P.F. Shall I grow sunflowers? Univ. of California Agr. Expt. Sta. Ext. Service Leaflet. Berkeley CA 

USA. [16 pages] 

1951 Knowles, P.F. and L.L. Davis. Safflower, a new oil crop. Univ. of California Agr. Expt. Sta. Ext. Service Leaflet. 


1953 Houston, B.R. and P.F. Knowles. Studies on Fusarium wilt of flax. Phytopath 43:491–495. 

1953 Knowles, P.F. and B.R. Houston. Resistance of flax varieties to Fusarium wilt. Agron J 45:408–414. 

1954 Knowles, P.F. and L.L. Davis. Do castor beans have a place? Univ. of California Agr. Expt. Sta. Ext. Service 

Leaflet. Berkeley CA USA. [14 pages] 

1954 Knowles, P.F. and L.L. Davis. Sunflowers as a field crop. Univ. of California Agr. Expt. Sta. Ext. Service Leaflet. 


1954 Knowles, P.F. and B.R. Houston. Inheritance of resistance to Fusarium wilt of flax in Dakota selection 

48–94. Agron J 47:131–135. 

1954 Knowles, P.F. and W.H. Lange. The sunflower moth: Preliminary experiments indicate parathion, DDT effective 

controls. Calif Agric 8(4):11–12. 

33

Appendix 2. continued 

1955 Knowles, P.F. Safflower—production, processing, and utilization. Econ Bot 9:273–299. 

1956 Knowles, P.F., B.R. Houston, and J.B. McOnie. Inheritance of resistance to Fusarium wilt of flax in Punjab 

53. Agron J 48:135–137. 

1957 Knowles, P.F., G.H. Abel, R.T. Edwards, and M.D. Miller. Soybean tips. Univ. of California Agr. Expt. Sta. 

Ext. Service Leaflet 94. Berkeley CA USA. 

1958 Knowles, P.F. Safflower. Adv Agron 10:289–323. 

1958 Knowles, P.F. and A. Ashri. Wild safflower in California: Improvement of cultivated safflower through plantbreeding 

program to obtain desirable characteristics of wild species. Calif Agric 12(4):4–5. 

1958 Knowles, P.F., R.T. Edwards, and M.D Miller. Soybeans in California. The Cotton Gin and Oil Mill Press 

March 8, 1958. 

1958 Zimmerman, L.H., M.D Miller, and P.F. Knowles. Castorbeans in California. Univ. of California Agr. Expt. 

Sta. Ext. Service Circular 468. Berkeley CA USA. [12 pages] 

1959 Ashri, A. and P.F. Knowles. Further notes on Carthamus in California. Leaflets in Western Bot 9:5–8. 

1959 Knowles, P.F. Plant Exploration report for safflower and miscellaneous oilseeds: Near East and Mediterranean 

countries, March-October 1958. Dept. of Agronomy, Univ. of California, Davis CA USA, in collaboration 

with USDA-ARS Crop Research Division, CR-43-5. [42 text pages plus photos] 

1959 Knowles, P.F. Oilseed crops in California. The Cotton Gin and Oil Mill Press October 1, 1959. 

1959 Knowles, P.F., W.H. Isom, and G.F. Worker. Flax production in the Imperial Valley. Univ. of California Agr. 

Expt. Sta. Ext. Service Circular 480. Berkeley CA USA. [28 pages] 

1959 Knowles, P.F., B.D. Sandlin, and M.D. Miller. Flax production in San Mateo County. Univ. of California Agr. 

Expt. Sta. Ext. Service Circular 482. Berkeley CA USA. [16 pages] 

1960 Ashri, A. and P.F. Knowles. Cytogenetics of safflower (Carthamus L.) species and their hybrids. Agron J 

52:11–17. 

1960 Knowles, P.F. New crop establishment. Econ Bot 14:263–275. 

1960 Knowles, P.F. Safflower’s native home. Crops and Soil 12:4. 

1960 Knowles, P.F. and M.D. Miller. Safflower in California. Univ. of California Agr. Expt. Sta. Ext. Service 

Manual 27. Berkeley CA USA. [26 pages] 

1960 Knowles, P.F. and M.D. Miller. Tips on safflower growing. Univ. of California Agr. Expt. Sta. Ext. Service 

Leaflet 126. Berkeley CA USA. [6 pages] 

1960 Yermanos, D.M. and P.F. Knowles. Effects of gibberellic acid treatment on safflower. Agron J 52:596–598. 

1961 Schank, S.C. and P.F. Knowles. Colchicine induced polyploids of Carthamus tinctorius L. Crop Sci 1:342– 

344. 

1962 Knowles, P.F. Complementary genes cause a lethal seedling character in flax. Crop Sci 2:270. 

1962 Knowles, P.F. Safflower oil mutant types under study. Calif Agric 16(4):7. 

1962 Yermanos, D.M. and P.F. Knowles. Fatty acid composition of the oil in crossed seed of flax. Crop Sci 2:109– 

111. 

1963 Knowles, P.F. Relationship and inheritance of fatty acids of safflower oil. in: Proc. Second Safflower Conference. 

Tucson AZ USA. 

1963 Knowles, P.F., M.D. Miller, and W.H. Isom. Safflower—An established crop in California. Univ. of California 

Agr. Expt. Sta. Ext. Service Leaflet 162. Berkeley CA USA. [12 pages] 

1963 Knowles, P.F. and A. Mutwakil. Inheritance of low iodine value of safflower selections from India. Econ Bot 

17:139–145. 

34


1964 Knowles, P.F. and A.B. Hill. Inheritance of fatty acid content in the seed oil of a safflower introduction from 

Iran. Crop Sci 4:406–409. 

1964 Knowles, P.F. and S.C. Schank. Artificial hybrids of Carthamus nitidus Boiss. and C. tinctorius L. (Compositae). 

Crop Sci 4:596–599. 

1964 Leon, R. and P.F. Knowles. Inheritance of appressed branching in safflower. Crop Sci 4:441. 

1964 Schank, S.C. and P.F. Knowles. Cytogenetics of hybrids of Carthamus species (Compositae) with ten pairs 

of chromosomes. Am J Bot 10:1903–1102. 

1965 Harvey, B.L. and P.F. Knowles. Natural and artificial alloploids with 22 pairs of chromosomes in the genus 

Carthamus (Compositae). Can J Genet Cytol 7:126–139. 

1965 Knowles, P.F. Oil plants. pages 902–904. Encyclopaedia Britannica. William Benton, Publisher. 

1965 Knowles, P.F. Variability in oleic and linoleic acid contents of safflower oil. Econ Bot 19:53–62. 

1965 Knowles, P.F. Report of Sabbatic Leave, August 1, 1964-August 1, 1965. Report for Univ. of California, 

Davis CA USA. [48 text pages plus photos] 

1965 Knowles, P.F., A.B. Bill, and J.E. Ruckman. High oleic acid content in new safflower, UC-1. Calif Agric 

19(12):15. [‘A.B. Bill’, as published, is actually ‘A.B. Hill’.] 

1965 Knowles, P.F. and M.D. Miller (with D.W. Henderson, Chester L. Foy, Elmer C. Carlson, J.M. Klisiewicz, 

John R. Goss, L.G. Jones, and Roydon T. Edwards). Safflower. Univ. of California Agr. Expt. Sta. Ext. Service 

Circular 532. Berkeley CA USA. 

1965 Worker, G.J. Jr., J.P. Jones, and P.F. Knowles. Safflower trials—Imperial Valley Field Station 1957 thru 1964. 

Field Crop Report No. 16. Dept. of Agronomy, Imperial Valley Field Station, Univ. of California Agr. Expt. 

Sta. Berkeley CA USA. [8 pages] 

1966 Ebert, W.W. and P.F. Knowles. Inheritance of pericarp types, sterility, and dwarfness in several safflower 

crosses. Crop Sci 6:579–582. 

1966 Knowles, P.F., A.B. Hill, and J.E. Ruckman. Safflower with unusual oil developed. Crops and Soil 18(5):19. 

1967 Briggs, F.N. and P.F. Knowles. Introduction to Plant Breeding. Reinhold Publishing Corp., New York NY 

USA. 

1967 Knowles, P.F. Processing seeds for oil in towns and villages of Turkey, India and Egypt. Econ Bot 21:156– 

162. 

1968 Ebert, W.W. and P.F. Knowles. Developmental and anatomical characteristics of thin-hull mutants of Carthamus 

tinctorius (Compositae). Am J Bot 55:421–430. 

1968 Hill, A.B. and P.F. Knowles. Fatty acid composition of the oil of developing seeds of different varieties of safflower. 

Crop Sci 8:275–277. 

1968 Knowles, P.F. Associations of high levels of oleic acid in the seed oil of safflower (Carthamus tinctorius) with 

other plant and seed characters. Econ Bot 22:195–200. 

1968 Knowles, P.F. Registration of UC-1 Safflower. (Reg. No. 6). Crop Sci 8:641. 

1968 Knowles, P.F., J.M. Klisiewicz, and A.B. Hill. Safflower introductions resistant to Fusarium wilt. Crop Sci 

8:636–637. 

1968 Knowles, P.F. and M.D. Miller (with D.W. Henderson, Chester L. Foy, Elmer C. Carlson, J.M. Klisiewicz, 

John R. Goss, L.G. Jones, and Roydon T. Edwards). El cártamo. (Univ. of California Agr. Expt. Sta. Ext. 

Service Circular 532. Berkeley CA USA). RTAC, USAID, USA. 

1969 Knowles, P.F. Centers of plant diversity and conservation of crop germ plasm: Safflower. Econ Bot 23:324– 

329. 

1969 Knowles, P.F. Modification of quantity and quality of safflower oil through plant breeding. J Am Oil Chem 

35


Soc 46:130–132. 

1969 Knowles, P.F. Breeding for changes in quality of safflower oil. pages 6–7 in: Proc. 39th Annual Flax Institute. 

1969 Knowles, P.F. Breeding safflower for changed oil composition. pages 25–26 in: P.F. Knowles and M.D. 

Miller (eds.) Proc. Third Safflower Research Conference, May 7–8, 1969, University of California, Davis. Univ. 

of California, Davis CA USA. 

1969 Knowles, P.F. Safflower. pages 302–303 in: McGraw-Hill Yearbook of Science and Technology. McGraw-Hill 

Publishing Company, New York NY USA. 

1969 Knowles, P.F. and M.D. Miller (eds.) Proc. Third Safflower Research Conference, May 7–8, 1969, University of 

California, Davis. Univ. of California, Davis CA USA. 

1969 G.O. Kohler, G. Fuller, R. Palter, and P.F. Knowles. Lysine content of safflower. pages 61–62 in: P.F. Knowles 

and M.D. Miller (eds.) Proc. Third Safflower Research Conference, May 7–8, 1969, University of California, 

Davis. Univ. of California, Davis CA USA. 

1969 Palter, R., G.O. Kohler, and P.F. Knowles. Survey for a high lysine variety in the world collection of safflower. 

Agric Food Chem 17:1298–1300. 

1970 Hockett, E.A. and P.F. Knowles. Inheritance of branching in sunflowers, Helianthus annuus L. Crop Sci 

10:432–436. 

1970 Imrie, B.C. and P.F. Knowles. Inheritance studies in interspecific hybrids between Carthamus flavescens and 

C. tinctorius. Crop Sci 10:349–352. 

1970 Khidir, M.O. and P.F. Knowles. Cytogenetic studies of Carthamus species (Compositae) with 32 pairs of 

chromosomes. I. Intrasectional hybridization. Am J Bot 57:123–129. 

1970 Khidir, M.O. and P.F. Knowles. Cytogenetic studies of Carthamus species (Compositae) with 32 pairs of 

chromosomes. II. Intersectional hybridization. Can J Genet Cytol 12:90–99. 

1970 Knowles, P.F., S.R. Temple, and F. Stolp. Variability in the fatty acid composition of sunflower seed oil. pages 

215–218 in: Proc. Fourth International Sunflower Conference, June 23–25, 1970. Memphis TN USA. 

1970 Ladd, S.L. and P.F. Knowles. Inheritance of stearic acid in the seed oil of safflower (Carthamus tinctorius L.). 

Crop Sci 10:525–527. 

1971 Beard, B.H. and P.F. Knowles. Frequency of cross pollination of soybeans after seed irradiation. Crop Sci 

11:681–684. 

1971 Imrie, B.C. and P.F. Knowles. Genetic studies of self-incompatibility in Carthamus flavescens Spreng. Crop 

Sci 11:489–492. 

1971 Knowles, P.F. Safflower introductions. FAO Plant Genetic Resources Newsl 25:19–21. 

1971 Ladd, S.L. and P.F. Knowles. Interactions of alleles at two loci regulating fatty acid composition of the seed 

oil of safflower (Carthamus tinctorius L.). Crop Sci 11:681–684. 

1972 Carlson, E.C., P.F. Knowles, and J.E. Dillé. Sunflower varietal resistance to sunflower moth larvae. Calif Agric 

26(6):11–13. 

1972 Knowles, P.F. The plant geneticist’s contribution toward changing lipid and amino acid composition of safflower. 

J Am Oil Chem Soc 49:27–29. 

1972 Urie, A.L. and P.F. Knowles. Safflower introductions resistant to Verticillium wilt. Crop Sci 12:545–546. 

1972 Worker, G.F. Jr. and P.F. Knowles. Safflower production under minimum and maximum soil preparation in 

Imperial Valley. Calif Agric 26(1):12–13. 

1973 Beard, B.H. and P.F. Knowles. Management practices for various areas in California. pages 18–20 in: B.H. 

Beard and P.F. Knowles (eds.) Soybean Research in California. Univ. of California Agr. Expt. Sta. Ext. Service 

Bull. 862. Berkeley CA USA. 

36


1973 Beard, B.H. and P.F. Knowles (eds.) Soybean Research in California. Univ. of California Agr. Expt. Sta. Ext. 

Service Bull. 862. Berkeley CA USA. [68 pages] 

1973 Beard, B.H. and P.F. Knowles. Varietal testing and improvement. pages 41–44 in: B.H. Beard and P.F. 

Knowles (eds.) Soybean Research in California. Univ. of California Agr. Expt. Sta. Ext. Service Bull. 862. 

Berkeley CA USA. 

1973 Beard, B.H., M.D. Miller, and P.F. Knowles. Introduction. pages 5–7 in: B.H. Beard and P.F. Knowles (eds.) 

Soybean Research in California. Univ. of California Agr. Expt. Sta. Ext. Service Bull. 862. Berkeley CA USA. 

1973 Knowles, P.F. Climate factors. pages 15–16 in: B.H. Beard and P.F. Knowles (eds.) Soybean Research in California. 

Univ. of California Agr. Expt. Sta. Ext. Service Bull. 862. Berkeley CA USA. 

1973 Knowles, P.F. Morphology and development of the soybean plant. pages 7–10 in: B.H. Beard and P.F. 

Knowles (eds.) Soybean Research in California. Univ. of California Agr. Expt. Sta. Ext. Service Bull. 862. 


1973 Knowles, P.F. New patterns in plant breeding. Editorial. Calif Agric 27(1):2. 

1973 Knowles, P.F., G.H. Abel, and B.H. Beard. Sowing methods and practices. pages 20–28 in: B.H. Beard and 

P.F. Knowles (eds.) Soybean Research in California. Univ. of California Agr. Expt. Sta. Ext. Service Bull. 862. 


1973 Knowles, P.F., D.B. Ferguson, and I.J. Johnson. Industry-University research assistantships. J Agron Educ 

2:89–90. 

1973 Lehman, W.F., F.E. Robinson, P.F. Knowles, and R.A. Flock. Sunflowers in the Desert Valley areas of Southern 

California. Calif Agric 27(8):12–14. 

1975 Ashri, A., D.E. Zimmer, A.L. Urie, and P.F. Knowles. Evaluation of the germ plasm collection of safflower 

Carthamus tinctorius L. VI. Length of planting to flowering period and plant height in Israel, Utah and Washington. 

Theor Appl Genet 46:359–364. 

1975 Dillé, J.E. and P.F. Knowles. Histology and inheritance of the closed flower in Carthamus tinctorius (Compositae). 

Am J Bot 62:209–215. 

1975 Knowles, P.F. Recent research on safflower, sunflower, and cotton. J Am Oil Chem Soc 52:374–376. 

1975 Temple, S.R. and P.F. Knowles. Inheritance of brittle stems in safflower. Crop Sci 15:694–697. 

1976 Beard, B.H. and P.F. Knowles. Improving protein supplies from oilseed crops and large-seeded legumes. 

pages 159–174 in: B.H. Beard and M.D. Miller (eds.) Opportunities to Improve Protein Quality and Quantity 

for Human Food. Univ. of California Spec. Publ. 3058. Davis CA USA. 

1976 Carapetian, J. and P.F. Knowles. Inheritance of genic sterility in cultivated safflower. Crop Sci 16:395–399. 

1976 Estilai, A. and P.F. Knowles. Cytogenetic studies of Carthamus divaricatus with eleven pairs of chromosomes 

and its relationship to other Carthamus species (Compositae). Am J Bot 63:771–782. 

1976 Knowles, P.F. Safflower. pages 31–33 in: N.W. Simmonds (ed.) Evolution of Crop Plants. Longman, London 

UK and New York NY USA. 

1976 Urie, A.L., L.H. Zimmerman, C.A. Thomas, W.F. Peterson, and P.F. Knowles. Registration of VFstp-1 safflower 

germplasm. Crop Sci 16:888. 

1977 Ashri, A., P.F. Knowles, A.L. Urie, D.E. Zimmer, A. Cahaner, and A. Marani. Evaluation of the germ plasm 

collection of Safflower, Carthamus tinctorius. III. Oil content and iodine value and their associations with 

other characters. Econ Bot 31:38–46. 

1977 Ghanavati, N.A. and P.F. Knowles. Variation among winter-type selections of safflower. Crop Sci 17:44–46. 

1977 Knowles, P.F. Safflower germplasm: Domesticated and wild. Calif Agric 31(9):12–13. 

1978 Estilai, A. and P.F. Knowles. Relationship of Carthamus leucocaulos to other Carthamus species (Composi- 

37


tae). Can J Genet Cytol 20:221–233. 

1978 Fernández-Martínez, J. and P.F. Knowles. Combined effects of genes for appressed and decumbent branching. 

Crop Sci 17:516–517. 

1978 Fernández-Martínez, J. and P.F. Knowles. Inheritance of self-incompatibility in wild sunflower, Helianthus 

annuus L. pages 484–489 in: Proc. 8th International Sunflower Conference, July 23–27, 1978. Minneapolis MN 

USA. 

1978 Heaton, T.C., P.F. Knowles, D.S. Mikkelsen, and J.E. Ruckman. Production of free fatty acids in safflower 

seeds by fungi. J Am Oil Chem Soc 55:465–468. 

1978 Khan, S.A. and P.F. Knowles. A programme for improving the quality of oil and meal in mustard and rape 

species of Pakistan. Pakistan Acad Sci 15:29–35. 

1978 Knowles, P.F. Morphology and anatomy. pages 55–87 in: J.F. Carter (ed.) Sunflower Science and Technology. 

Am. Soc. Agron. Monograph, Madison WI USA. 

1979 Knowles, P.F. Introduction, promotion and production of soybeans as an irrigated crop. pages 9–15 in: 

Prelim. Proc. Conference on Irrigated Soybean Production in Arid and Semi-Arid Regions. Cairo, EGYPT. 

1979 Knowles, P.F. and T.E. Kearney. Rapeseed production in California. Agronomy Progress Report no. 102. 

Univ. of California Cooperative Extension Service, Berkeley CA USA. 

1979 Urie, A.L., W.F. Peterson, and P.F. Knowles. Registration of Oleic Leed safflower (Reg. No. 8). Crop Sci 

19:747. 

1980 Estilai, A. and P.F. Knowles. Aneuploids in safflower. Crop Sci 20:516–518. 

1980 Heaton, T.C. and P.F. Knowles. Registration of UC-148 and UC-149 male-sterile safflower germplasm. 

(Reg. Nos. GP-16 and GP-17). Crop Sci 20:554. 

1980 Knowles, P.F. Safflower. pages 535–548 in: W.R. Fehr and H.H. Hadley (eds.) Hybridization of Crop Plants. 

Am. Soc. Agron. - Crop Sci Soc. Am., Madison WI USA. 

1980 Knowles, P.F. and R. Bukantis. Safflower. pages 131–133 in: D. Pimentel (eds.) Handbook of Energy Utilization 

in Agriculture. CRC Press, Boca Raton FL USA. 

1980 Smith, D., G. Fick, F. Johnson, and P.F. Knowles. Seed set in sunflower. The Sunflower 6(5):10–12. 

1980 Urie, A.L., D.J. DaVia, P.F. Knowles, and L.H. Zimmerman. Registration of 14-5 safflower germplasm (Reg. 

No. GP 15). Crop Sci 20:115–116. 

1981 DaVia, D.J., P.F. Knowles, and J.M. Klisiewicz. Evaluation of the World Safflower Collection for resistance to 

Phytophthora. Crop Sci 21:226–229. 

1981 Futehally, S. and P.F. Knowles. Inheritance of very high levels of linoleic acid in an introduction of safflower 

(Carthamus tinctorius L.) from Portugal. pages 56–61 in: P.F. Knowles (ed.) Proc. First International Safflower 

Conference, Davis, California, July 12–16, 1981. Univ. of California, Davis CA USA. 

1981 George, D.L., P.F. Knowles, and S.E. Shein. Compatibility, autogamy and environmental effects on seedset 

in selected sunflower hybrids and their inbred parents. pages 140–146 in: Proc. 9th International Conference 

of Sunflower, Torremolinos, Malaga (Spain), 8-13 June 1980. Ministerio de Agricultura, Servicio de Publicaciones 

Agrarias, Madrid SPAIN. 

1981 Knowles, P.F. (ed.). Proc. First International Safflower Conference. Univ. of California, Davis CA USA. July 

12–16, 1981. Univ. of California, Davis CA USA. [299 pages] 

1981 Knowles, P.F., T.E. Kearney, and D.B. Cohen. Species of rapeseed and mustard as oil crops in California. 

pages 255–268 in: E.H. Pryde, L.H. Princen, and K.D. Mukherjee (eds.). New Sources of Fats and Oils. 

Monogr. No. 9, Am. Oil Chem. Soc., Champaign IL USA. 

1982 Fernández-Martínez, J. and P.F. Knowles. Genética de la polifloria en el girasol silvestre Helianthus annuus L. 

Anales del Instituto Nacional de Investigaciones Agrarias Serie: Agricola 17:25–30. 

38


1982 Fernández-Martínez, J. and P.F. Knowles. Maternal and embryo effects on the oleic and linoleic acid contents 

of sunflower oil. pages 241–243 in: Proc. 10th International Sunflower Conf. Surfers Paradise, Australia, 

March 14–18, 1982. Australian Sunflower Association. Toowoomba QLD AUSTRALIA. 

1982 George, D.L., S.E. Shein, and P.F. Knowles. Effect of isolation bags on seedset in sunflower. pages 224–225 

in: Proc. 10th International Sunflower Conf. Surfers Paradise, Australia, March 14–18, 1982. Australian Sunflower 

Association. Toowoomba QLD AUSTRALIA. 

1982 George, D.L., S.E. Shein, and P.F. Knowles. Effect of stigmatic manipulation on pollination and seedset 

in sunflower. pages 226–227 in: Proc. 10th International Sunflower Conf. Surfers Paradise, Australia, March 

14–18, 1982. Australian Sunflower Association. Toowoomba QLD AUSTRALIA. 

1982 Klisiewicz, J.M. and A.L. Urie. Registration of Fusarium resistant safflower germplasm. Crop Sci 22:165. 

1982 Knowles, P.F. Safflower: Genetics and breeding. pages 89–101 in: Anon. (ed.) Improvement of Oil-Seed and 

Industrial Crops by Induced Mutations. International Atomic Energy Agency, Vienna AUSTRIA. 

1982 Knowles, P.F. 2. Plant breeding: Successes and limitations. Calif Agric 36(8):6–7. 

1983 Cohen, D.B. and P.F. Knowles. Evaluation of Brassica species in California. pages 282–287 in: Proc., 6th 

International Rapeseed Conference, Paris, May 1983, v. 1. Groupe Consultatif International de Recherche sur 

le Colza. Paris FRANCE. 

1983 Knowles, P.F. Genetics and breeding of oilseed crops. Econ Bot 37:423–433. 

1983 Lyon, C.K., P.F. Knowles, and G.O. Kohler. Evaluation of Brassica species as leaf sources for extending the 

processing season of a leaf protein concentrate plant. J Sci Food Agric 34:849–854. 

1983 Sailsbery, R.L. and P.F. Knowles. Hybrid Sunflower Seed Production. Publ. Coop. Extn., Glenn County, Univ. 

of California, Berkeley CA USA. 

1983 Tysdal, H.M., A. Estilai, I.A. Siddiqui, and P.F. Knowles. Registration of four guayule germplasms. Crop Sci 

23:189. 

1984 Hirsinger, F. and P.F. Knowles. Morphological and agronomic description of selected Cuphea germplasm 

[Mexico]. Econ Bot 38:439–451. 

1985 Knowles, P.F. Oilseed crops breeding: Past and future. pages 199–209 in: A. Omran (ed.) Oil Crops: Sesame 

and Safflower. Proceedings of the Second Oil Crops Network Workshop Held in Hyderabad, India, 5–9 

February 1985. (Manuscript Report, IDRC-MR105e). 

1988 Knowles, P.F. Recent advances in oil crops breeding. pages 35–38 in: T.H. Applewhite (ed.) Proc.: World 

Conference on Biotechnology for the Fats and Oils Industry. Am. Oil Chem. Soc., Champaign IL USA. 

1989 Knowles, P.F. Genetics and breeding of oil crops. pages 260–282 in: G. Röbbelen, R.K. Downey, and A. 

Ashri (eds.) Oil Crops of the World. McGraw Hill, New York NY USA. 

1989 Knowles, P.F. Safflower. pages 363–374 in: G. Röbbelen, R.K. Downey, and A. Ashri (eds.) Oil Crops of the 

World. McGraw Hill, New York NY USA. 

1989 Knowles, P.F. Safflower in China: A report of a visit made in 1988. Report for Univ. of California, Davis CA 

USA. [50 pages] 

1991 Knowles, P.F. Global perspectives of safflower. pages 13–16 in: V. Ranga Rao and M. Ramachandran, (eds.) 

Proc. Second International Safflower Conference, Hyderabad, India, 9–13 January 1989. Indian Society of Oilseeds 

Research, Directorate of Oilseeds Research, Hyderabad INDIA. 

1993 Carapetian, J. and P.F. Knowles. Genetic linkage between the trigenic male-female sterility and oil quality 

alleles in safflower. Crop Sci 33:239–242. 

1995 Knowles, P.F. and A. Ashri. Safflower, Carthamus tinctorius (Compositae). pages 47–50 in: J. Smartt and 

N.W. Simmonds (eds.) Evolution of Crop Plants, 2nd Ed. Longman Scientific & Technical. Essex UK and J. 

Wiley and Sons, NY USA. 

39

Appendix 3. List by species, year, and PI number 

of 1175 safflower accessions associated with P.F. 

Knowles in the collection at the USDA-ARS-RPIS, 

Pullman WA USA 1 . 

1 Editors’ note: List was current as of a 1995 request to the USDA Pullman Regional Plant Introduction Station (and includes an additional 93 accessions 

from a 1988 safflower nursery at the Beijing Botanical Garden selected by PF Knowles for acquisition by the RPIS). Current status of any accession can be 

determined via query by PI number at the USDA Genetic Resources Information Network (http://www.ars-grin.gov/npgs/acc/acc_queries.html). 

Carthamus tinctorius 

1958 248355, 248356, 248357, 248358, 248359, 248360, 248361, 248362, 248363, 248364, 248365, 

248366, 248367, 248368, 248369, 248370, 248371, 248372, 248373, 248374, 248375, 248376, 

248377, 248378, 248379, 248380, 248381, 248382, 248383, 248384, 248385, 248386, 248387, 

248388, 248389, 248624, 248625, 248626, 248627, 248628, 248629, 248630, 248631, 248632, 

248633, 248793, 248794, 248795, 248796, 248797, 248798, 248799, 248800, 248801, 248802, 

248803, 248804, 248805, 248806, 248807, 248808, 248809, 248810, 248811, 248812, 248813, 

248814, 248815, 248816, 248817, 248818, 248819, 248820, 248821, 248822, 248823, 248824, 

248825, 248826, 248827, 248828, 248829, 248830, 248831, 248832, 248833, 248834, 248835, 

248836, 248837, 248838, 248839, 248840, 248841, 248842, 248843, 248844, 248845, 248846, 

248847, 248848, 248849, 248850, 248851, 248852, 248853, 248854, 248855, 248856, 248857, 

248858, 248859, 248860, 248861, 248862, 248863, 248864, 248865, 248866, 248867, 248868, 

248869, 248870, 248871, 248872, 248873, 248874, 248875, 248876, 248877, 248878, 248879, 

248880, 250006, 250007, 250008, 250009, 250010, 250011, 250075, 250076, 250077, 250078, 

250079, 250080, 250081, 250082, 250083, 250179, 250180, 250182, 250183, 250186, 250187, 

250188, 250189, 250190, 250191, 250192, 250193, 250194, 250195, 250196, 250197, 250198, 

250199, 250200, 250201, 250202, 250203, 250204, 250205, 250208, 250335, 250336, 250337, 

250338, 250341, 250342, 250343, 250345, 250346, 250347, 250348, 250350, 250351, 250353, 

250473, 250474, 250475, 250476, 250477, 250478, 250479, 250480, 250481, 250482, 250523, 

250524, 250525, 250526, 250527, 250528, 250529, 250530, 250531, 250532, 250533, 250534, 

41



1958 continued 

250535, 250536, 250537, 250538, 250539, 250540, 250541, 250595, 250596, 250597, 250598, 

250599, 250600, 250601, 250603, 250604, 250605, 250606, 250607, 250608, 250609, 250610, 

250611, 250708, 250709, 250710, 250711, 250712, 250713, 250714, 250715, 250716, 250717, 

250718, 250719, 250720, 250721, 250722, 250723, 250724, 250819, 250820, 250821, 250822, 

250823, 250824, 250825, 250826, 250827, 250828, 250829, 250830, 250831, 250832, 250833, 

250834, 250835, 250836, 250837, 250838, 250839, 250840, 250841, 250842, 250920, 250921, 

250922, 250924, 250925, 250926, 251262, 251264, 251265, 251266, 251267, 251268, 251284, 

251285, 251288, 251289, 251290, 251291, 251398, 251462, 251977, 251978, 251979, 251980, 

251981, 251982, 251983, 251984, 251985, 251986, 251987, 251988, 251989, 252040, 252041, 

252042, 253384, 253385, 253386, 253387, 253388, 253389, 253390, 253391, 253392, 253393, 

253394, 253395, 253396, 253511, 253512, 253513, 253515, 253516, 253517, 253518, 253519, 

253520, 253521, 253522, 253523, 253524, 253527, 253528, 253529, 253531, 253534, 253535, 

253537, 253538, 253540, 253541, 253542, 253543, 253544, 253546, 253547, 253548, 253549, 

253550, 253552, 253553, 253554, 253555, 253556, 253559, 253560, 253561, 253562, 253563, 

253564, 253566, 253567, 253568, 253569, 253570, 253571, 253758, 253759, 253761, 253762, 

253763, 253764, 253765, 253892, 253893, 253894, 253895, 253896, 253897, 253898, 253899, 

253900, 253902, 253903, 253905, 253906, 253907, 253908, 253909, 253910, 253911, 253912, 

253913, 253914, 253916, 254090 

1959 254976 

1964 301047, 301048, 301049, 301050, 301051, 301052, 301053, 301054, 301055 

1965 304408, 304409, 304411, 304437, 304438, 304439, 304440, 304441, 304442, 304443, 304444, 

304445, 304446, 304447, 304448, 304449, 304450, 304451, 304452, 304453, 304454, 304455, 

304456, 304457, 304458, 304459, 304460, 304461, 304462, 304463, 304464, 304465, 304466, 

304467, 304468, 304469, 304470, 304471, 304472, 304473, 304474, 304475, 304476, 304494, 

304495, 304496, 304497, 304498, 304499, 304500, 304501, 304502, 304503, 304504, 304505, 

304506, 304507, 304508, 304509, 304510, 304590, 304591, 304592, 304593, 304594, 304595, 

304596, 304597, 304598, 304599, 305151, 305152, 305153, 305154, 305155, 305156, 305157, 

305159, 305160, 305161, 305162, 305163, 305164, 305165, 305166, 305167, 305168, 305169, 

305170, 305171, 305173, 305174, 305175, 305176, 305178, 305179, 305181, 305182, 305185, 

305186, 305187, 305188, 305189, 305190, 305191, 305192, 305193, 305195, 305196, 305197, 

305198, 305199, 305200, 305202, 305204, 305205, 305206, 305207, 305208, 305209, 305210, 

305211, 305212, 305214, 305215, 305216, 305217, 305218, 305219, 305220, 305221, 305253, 

306593, 306594, 306595, 306596, 306597, 306598, 306599, 306600, 306601, 306602, 306603, 

306604, 306605, 306606, 306607, 306608, 306609, 306610, 306611, 306612, 306613, 306614, 

306820, 306821, 306822, 306823, 306824, 306825, 306826, 306827, 306828, 306829, 306830, 

306831, 306832, 306833, 306836, 306838, 306839, 306840, 306841, 306842, 306843, 306844, 

306845, 306846, 306847, 306848, 306849, 306850, 306851, 306852, 306853, 306854, 306855, 

306856, 306857, 306858, 306861, 306865, 306866, 306867, 306868, 306869, 306870, 306872, 

306873, 306874, 306875, 306876, 306877, 306878, 306879, 306880, 306881, 306883, 306885, 

306886, 306887, 306888, 306889, 306890, 306891, 306892, 306894, 306896, 306897, 306898, 

306899, 306900, 306901, 306902, 306903, 306904, 306905, 306906, 306907, 306908, 306909, 

306910, 306911, 306912, 306913, 306915, 306916, 306917, 306918, 306920, 306921, 306922, 

306923, 306924, 306925, 306926, 306927, 306928, 306929, 306930, 306931, 306932, 306933, 

306934, 306935, 306936, 306939, 306940, 306941, 306942, 306943, 306944, 306945, 306946, 

306947, 306948, 306949, 306950, 306951, 306952, 306953, 306954, 306955, 306956, 306957, 

306958, 306959, 306960, 306961, 306962, 306963, 306964, 306965, 306967, 306968, 306969, 

306970, 306971, 306972, 306973, 306974, 306975, 306976, 306977, 306978, 306979, 306980, 

42



1965 continued 

306981, 306983, 306984, 306985, 306986, 306987, 306988, 306989, 306991, 306992, 306993, 

306994, 306995, 306996, 306997, 306998, 306999, 307000, 307001, 307002, 307003, 307004, 

307005, 307006, 307007, 307008, 307009, 307010, 307011, 307012, 307013, 307014, 307015, 

307016, 307018, 307019, 307020, 307021, 307022, 307023, 307024, 307025, 307026, 307027, 

307028, 307029, 307030, 307031, 307032, 307033, 307034, 307035, 307036, 307037, 307038, 

307039, 307040, 307041, 307042, 307043, 307044, 307045, 307046, 307047, 307048, 307049, 

307050, 307051, 307052, 307053, 307054, 307055, 307056, 307057, 307058, 307059, 307060, 

307061, 307062, 307063, 307064, 307065, 307066, 307067, 307068, 307069, 307070, 307071, 

307072, 307073, 307074, 307075, 307076, 307077, 307078, 307079, 307080, 307081, 307082, 

307083, 307084, 307085, 307086, 307087, 307088, 307089, 307090, 307091, 307092, 307093, 

307094, 307095, 307096, 307097, 307098, 307099, 307100, 307101, 307102, 307103, 307104, 

307105, 307106, 307107, 307108, 307109, 307110, 307111, 307112, 307113, 307114, 307115, 

307116, 307117, 307118, 307119, 307120, 307121, 307122, 307123, 307124, 307125, 307126, 

307127, 307128, 307129, 307130, 307131, 307132, 307133, 307134, 307135, 307136, 307137, 

307138 

1967 572434 

1969 343772, 343773, 343774, 343775, 343776, 343777, 343778, 343779, 343780, 343781, 343782, 

343783 

1972 372914 

1974 387820, 387821, 393498, 393499, 393500 

1975 405954, 405955, 405956, 405957, 405958, 405959, 405960, 405961, 405962, 405963, 405964, 

405965, 405966, 405967, 405968, 405969, 405970, 405971, 405972, 405973, 405974, 405975, 

405976, 405977, 405978, 405979, 405980, 405981, 405982, 405983, 405984, 405985, 405986, 

405987, 405988, 405989, 405990, 405991, 405992, 405993, 405994, 405995, 405996, 405997, 

405998, 405999, 406000, 406001, 406002, 406003, 406004, 406005, 406006, 406007, 406008, 

406009, 406010, 406011, 406012, 406013, 406014, 406015, 406016, 406017, 406018, 406019, 

406020, 407605, 407606, 407607, 407608, 407609, 407610, 407611, 407612, 407613, 407614, 

407615, 407616, 407617, 407618, 407619, 407620, 407621, 407622, 407623, 407624 

1978 426186, 426521, 426523, 426524 

1989 543974, 543975, 543976, 543977, 543978, 543979, 543980, 543981, 543982, 543983, 543984, 

543985, 543986, 543987, 543988, 543989, 543990, 543991, 543992, 543993, 543994, 543995, 

543996, 543997, 543998, 543999, 544000, 544001, 544002, 544003, 544004, 544005, 544008, 

544009, 544010, 544011, 544012, 544013, 544014, 544015, 544016, 544017, 544018, 544019, 

544021, 544022, 544023, 544024, 544025, 544026, 544027, 544028, 544029, 544030, 544031, 

544032, 544033, 544034, 544035, 544036, 544037, 544038, 544039, 544040, 544041, 544042, 

544043, 544044, 544045, 544046, 544047, 544048, 544049, 544050, 544051, 544052, 544053, 

544054, 544055, 544056, 544057, 544058, 544059, 544060, 544061, 544062, 576979, 576980, 

576981, 576982, 576983, 576984, 576985, 576986, 576987, 576988, 576989, 653133, 653134, 

653135, 657784 

1991 544063, 560161, 560162, 560176, 560177 

43


Carthamus oxyacantha 

1958 249970, 252022, 253843 

1975 407602 

1978 426182, 426183, 426184, 426185, 426427, 426428, 426429, 426430, 426431, 426433, 426434, 

426435, 426436, 426437, 426438, 426439, 426440, 426441, 426442, 426443, 426444, 426445, 

426447, 426449, 426450, 426451, 426452, 426454, 426456, 426457, 426458, 426459, 426463, 

426464, 426467, 426468, 426469, 426470, 426472, 426473, 426476, 426477, 426478, 426479, 

426480, 426481, 426482, 426483, 426484, 426485, 426486, 426487, 426488, 426489, 426490, 

426492, 426493, 426494, 426495, 426496, 426498, 426499, 426500, 426501, 426502, 426503, 

426504, 426505, 426506, 426507, 426508, 426509, 426510, 426511, 426512, 426514, 426515, 

426516, 426517, 426518, 426519, 426520 

Carthamus glaucus 

1958 2551254 

Carthamus glaucus alexandrinus 

1958 250588 

Carthamus lanatus 

1958 2553862 

1989 544020 

Carthamus lanatus turkestanicus 

1978 426180, 426181, 426425 ,426426 

Carthamus sp. 

1978 426187, 426188, 426189 

44

Safflower in California - University of California Cooperative Extension

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