Proceedings of the Iowa Academy of Science

Volume 85 Number Article 4

1978

Morphological and Cytological Relationships Between robusta and Stipa viridula (Gramineae: )

Margarita A. Mondrus Iowa State University

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Recommended Citation Mondrus, Margarita A. (1978) "Morphological and Cytological Relationships Between Stipa robusta and Stipa viridula (Gramineae: Stipeae)," Proceedings of the Iowa Academy of Science, 85(3), 84-87. Available at: https://scholarworks.uni.edu/pias/vol85/iss3/4

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Proc. Iowa Acad. Sci. 85(3): 84-87. 1978

Morphological and Cytological Relationships Between Stipa robusta and Stipa viridula (Gramineae: Stipeae)1

MARGARITA A. MONDRUS2

(Department of Botany and Pathology, Iowa State University, Ames, Iowa 50013).

Stipa robusta (Vasey) Scribn. andStipa viridula Trin. have previously been distinguished on the basis of overlapping quantitative characters, which often are inadequate for identification. Recent chromosome counts had not been made for either species. I have distinguished several effective diagnostic characters, including palea length, shape and pubescence, ratio of lemma length to width, ratio of palea length to lemma length, and panicle density. The most reliable character is pal ea length. INDEX DESCRIPTORS: Gramineae, , Stipeae, Stipa, forages, range.

Stipa viridula Trin. and S. robusta (Vasey) Scribn. are perennial Mitotic chromosome counts were obtained by a technique modified caespitose grasses of the Plains and Rocky Mountain regions. Stipa from that described by Sun and Sadanaga ( 1968) for root tip counts of viridula was first described by Trinius ( 1838) from the type specimen. seedlings of sativa. Root tips were prefixed in a saturated Vasey ( 1892) distinguished S. viridula var. robusta on the basis of its solution of paradichlorobenzene at 15°C for 3 hr and fixed in 3: 1 generally larger size. Vasey ( 1887) noted that S. robusta has a toxic absolute ethanol and glacial acetic acid at 37°C for 48 hr, hydrolyzed in effect on livestock. This chemical property, not found in S. viridula, IN HCI, and transferred to Schiffs Reagent for staining. Pectinase which is a palatable forage grass, has given S. robusta the common was used to dissolve the middle lamellae, and root tips were squashed in name of'' Sleepy Grass.'' Stipa robusta was raised to the species level propio-carmine. Photographs of chromosomes were taken with a Nikon by Lamson-Scribner ( 1897), who changed the name toStipa vaseyi the 4 x 5 plate camera. following year (Lamson-Scribner, 1898) because of the previous use of S. robusta Nutt. as a synonym of S. spartea Trin. (Trinius and OBSERVATIONS Ruprecht, 1849). Hitchcock (1935) changed the name back to S. robusta (Vasey) Scribn. Paleas of Stipa viridula range in length from 1.0 to 2.5 mm long and Hitchcock ( 1951) distinguished S. viridula and S. robust a on the are elliptical 2. 9: I, hyaline, and have acute glabrous apices. Paleas of basis of glume texture, plant height, and panicle density. He did not Stipa robusta are firm, 3.5 to 5.0 mm long, oblong 5.3: I, with truncate, report any differences in palea or lemma characters. Distinctions ciliate apices. based on morphological characters have not been sharply delineated Lemmas of S. viridula are mostly 4.5 to 6.5 mm long and I to 1.5 by previous investigators. mm wide at maturity. The ratio of lemma length to width is mostly 5: 1 Small chromosomes are characteristic of the Stipeae (Stebbins, or less. Mature lemmas of S. robusta range from 6.0 to 8.0 mm in 1956). A basic chromosome number of n=6 has been suggested for length and from 0.8 to 1.2 mm in width. The ratio of lemma length to Stipa (Johnson, 1972). Johnson and Rogler (1943) determined the width is mostly greater than 5: 1. Glumes of S. viridula usually are somatic chromosome number of S. viridula as 2n =82. A mitotic hyaline, whereas those of S. robusta are firm. chromosome count of 2n=64 for S. robusta was previously deter­ Panicles of S. viridula are I 0 to 30 cm long, narrow, and somewhat mined by Love (Myers, 1947). I have confirmed both these previous compressed, with an average of 10 to 15 spikelets per cm2 . Panicles of counts. S. robusta are similar in length, more compressed than those of S. In this paper, I have clarified the interrelationships and delimitation viridula, and more densely-flowered, with an average of 15 to 25 of these two entities and have proposed a taxonomic treatment based spikelets per cm2 . on my observations. I determined a somatic chromosome number of 2n = 82 for S. vir-

MATERIALS AND METHODS Table 1. Variation in selected characters of S. viridula and S. robusta Herbarium specimens, mass collections, and immature inflores­ cences were collected throughout the Plains and Rocky Mountain Character S. viridula S. robusta states. Herbarium specimens from the following herbaria were Panicle density (6) 10-15 (17) examined: Ariz., ISC, MO, NY, US. Spikelets soaked .in a softening (8) 15-25 (28) in spikelets/cm2 Mean=8.39 Mean=l5.59 solution (Pohl, 1965) were measured with a Bausch and Lomb dissect­ ing microscope fitted with an ocular micrometer disc with 0.1-mm Plant height (31) 40-115 (148) (42) 60-160 (178) divisions. Shapes and ratios of floral parts are given according to the in cm Mean=78.02 Mean=89.73 Systematics Association Committee for Descriptive Biological Ter­ Length of lemma (4.0) 4.5-6.5 (8.2) (5.5) 6.0-8.0 (8.8) minology (1962). Panicle density was determined by counting the in mm Mean=5.33 Mean=6.83 number of spikelets visible through a l-cm2 frame. Width of lemma (0.6) 1.0-1.5 (1.8) (0.6) 0.8-1.2 ( 1.5) in mm Mean= 1.05 Mean=l.01 Lemma length to (2.8) 3.5-6.5 (10.0): 1 (4.4) 5.0-8.0 (12.0): l 'The facilities of the Iowa State University Herbarium, supported by the Sci­ width ratio Mean=5.05: l Mean=6.90:1 ences and Humanities Research Institute, were used in the preparation of this Length of palea (0.8) 1.0-2.5 (3.5) (3.0) 3.5-5.0 (6.0) paper. Journal Paper No. J-9077 of the Iowa Agriculture and Home Economics in mm Mean= 1.99 Mean=4.39 Experiment Station, Ames, Iowa 50011; Project 1833. Ratio of palea (0.20) 0.22-0.38 (0.43) (0.55) 0.58-0.63 (0.68) 2 Current address: Plant and Soil Science Dept., State Polytechnic length to lemma Mean=0.37 Mean=0.64 University, Pomona, California 91768. length

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MORPHOLOGY AND CYTOLOGY OF TWO ST/PA SPECIES 85

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• • • • • • • • • 1 2 •

Fig. I. Mitotic chromosomes of Stipa viridula; 2n =82; 2847 x Fig. 2. Mitotic chromosomes of Stipa robusta; 2n=64; 2847 X

idula. A photograph of the chromosomes is shown in Fig. I. The source of these species, pal ea length has been found diagnostic for the Chilean of the somatic material was Mondrus 122 collected in Pueblo County, species of Stipa (Matthei, 1965) and for the California species of Stipa . (Dedecca, 1954). The shape of the floret also is diagnostic, as shown in I determined a somatic chromosome number of 2n = 64 for S. Fig. 4. This character is expressed in this paper in terms of the ratio of robusta. The chromosomes are shown in Fig. 2. The material was from length to width of the intact lemma. Although there is some overl ap of Mondrus 173 collected in Lincoln County, . Voucher the ranges of variation of lemma length and width and little difference specimens for both counts are deposited in the Iowa State University between the means of either length or width alone, the difference Herbarium. becomes evident when shape of floret is expressed in terms of the ratio of length to width. The mean values are notably different. Various DISCUSSION ratios of floral parts have been found diagnostic for certain species of Stipa and by Johnson ( 1945, 1962), Dedecca ( 1954), and Stipa viridula and Stipa robusta are caespitose perennials very simi­ Schechter and Johnson ( 1966). lar in external morphology. Vegetative characters are inadequate for The texture of the glumes is another distinguishing character. The diagnostic purposes. Plant height has traditionally been used as a major glumes of S. viridula are almost always hyaline. The glumes of S. distinguishing character. I found considerable overl ap of the ranges of robusta are firm. A few intermediate specimens were observed. variation of plant height. From Table I it is evident that plant height is Figure 3 shows data from herbarium specimens and my own collec­ not a reliable diagnostic character. Other vegetative characters also are tions, representing the entire geographic range of each species, plotted highly variable. on a scatter diagram. There is an obvious clustering of specimens, Panic le density and spikelet characters are highly diagnostic. Table I because of the difference in palea length. Plant height shows a con­ gives the ranges, extremes, and mean values for these characters, tinuum, with S. viridula at the low end and S. robusta at the high end . which are shown pictorially in Fig. 3. The length of the palea probably Panicle density , glume texture, and lemma length-to-width ratio also is the single most diagnostic character. There is virtually no overlap of are shown to be diagnostic. the ranges of variation, and only occasionally are specimens inter­ Because of the significant differences in chromosome number, mediate between the high value for S. viridula and the low value for S. spikelet morphology, and panicle density , it is recommended that the robusta. Although I have seen no keys that mention the palea of either two taxa continue to be treated as separate species. Palea characters and https://scholarworks.uni.edu/pias/vol85/iss3/4 2 Mondrus: Morphological and Cytological Relationships Between Stipa robusta

86 PROC. IOWA ACAD. SCI. 85 (1978)

60

5.5

50

45

40

-<) ~ 0- ()-- 0--

-0- -0 GLUMES 2.0 0 () • hyoline _. 1ndurate 0- LEMMA 1.5 length to Width Ratio -D-0 0 >10 510 <5

l.O Den~1tf~~~~~~s cm2 0-0-0 < 10 10-15 > 15 0.5 3 30 40 50 60 70 80 90 110 120 130 140 150 160 170 180 PLANT HEIGHT IN CM.

Fig. 3. Scatter diagram of 75 herbarium specimens of Stipa viridula and Stipa robusta.

ratios of floral parts are effective key characters that have not been used JOHNSON, B. L. 1945. Cyto-taxonomic studies in Oryzopsis. Bot. Gaz. 107: ·previously to distinguish these species. Use of these characters in keys 1-32. would facilitate identification of Stipa viridula and Stipa robust a. JOHNSON, B. L. 1962. Natural hybrids between Oryzopsis and Stipa: II. X Stipa nevadensis. Am. J. Bot. 49: 540-546. JOHNSON, B. L. 1972. Polyploidy as a factor in the evolution and distribution ACKNOWLEDGMENTS of grasses. Pages 18-35 in V. B. Youngner, ed. The biology and utilization of grasses. Academic Press, New York. I sincerely thank Dr. Richard Pohl for his support, advice, and JOHNSON, B. L. and G. A. ROGLER. 1943. A cyto-taxonomic study of an helpful criticism throughout this study. Special thanks are extended to intergeneric hybrid between Oryzopsis hymenoides andStipa viridu/a. Am. Holly S. Heer of the Iowa State University Soybean Cytogenetics J. Bot. 30: 49-56. Laboratory for her assistance in preparation of root tips for mitotic LAMSON-SCRIBNER, F. 1897. Page 23 in P.A. Rydberg and C. L. Shear. A chromosome counts and for the photographs of chromosomes. I am report upon the grasses and forage of the Rocky Mountain region. grateful to the curators who loaned specimens of S. viridula and S. U.S. Dep. Agric. Div. Agrost. Bull. 5: 1-45. LAMSON-SCRIBNER, F. 1898. Studies on American grasses: II. Descriptions robust a for this study. of little known grasses. U.S. Dep. Agric. Div. Agrost. Bull. 11: 42-58. MATIHEI, 0. R. 1965. Estudio critico de las gramineas de! genero Stipa en Chile. Gayana Bot. 13: 1-136. MYERS, W. M. 1947. Cytology and genetics of forage grasses. Bot. Rev. 13: LITERATURE CITED 319-421. POHL. R. W. 1965. Dissecting equipment and materials for the study of minute DEDECCA, D. M. 1954. Studies on the California species of Stipa plant structures. Rhodora 67: 95-96. (Gramineae). Madrono 12: 129-139. SCHECHTER, Y., and B. L. JOHNSON. 1966. A new species ofOryzopsis HITCHCOCK, A. S. 1935. Manual of the grasses of the United States; U.S. (Gramineae) from . Brittonia 18: 342-347. Dep. Agric. Misc. Pub!. 200. STEBBINS, G. L. 1956. Cytogenetics and evolution of the grass family. Am. J. HITCHCOCK, A. S. 1951. Manual of the grasses of the United States. 2nd ed. Bot. 43: 890-905. Revised by A. Chase. U.S. Dep. Agric. Misc. Pub!. 200. SUN, N., AND K. SADANAGA. 1968. Handling of oat chromosomes. Crop.

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MORPHOLOGY AND CYTOLOGY OF TWO ST/PA SPECIES 87

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Fig. 4. Floret, lemma and palea of Stipa viridula (left); Floret, lemma and palea of Stipa robusta (right)

Sci. 8: 767-769. TRINIUS, C. B., and F. J. RUPRECHT. 1849. Gramina agrostidea. III. Callus SYSTEMATICS ASSOCIATION COMMITTEE FOR DESCRIPTIVE obconicus (Stipacea). Mem. Acad. St. Petersb. 7(2): 1-189. BIOLOGICAL TERMINOLOGY. 1962. Terminology of simple symmetri­ VASEY, G. 1887. Special uses and properties of some Mexican grasses. Bull. cal plane shapes. Taxon II: 145-156. Torrey Bot. Club 14: 98-100. TRINIUS, C. B. 1838. Graminum suppl. Mem. Acad. St. Petersb. Ser. VI, VASEY, G. 1892. Monograph of the grasses of the United States and British 4(2): 1-107. America. Contrib. U.S. Natl. Herb. 3: 1-89.

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