2 8 2 5 2 5 1.0 :: 11111 . 11111 . 1.0 :; IllFa 11111 . ~ OOI3.~ !i,g 1~1I3.2 W 2.2 I:J a.:;. E~ a.:;. I~ w ~ &:0; I!i 2.0 '­ e~ &.:: B~ ... ~ ... " I 1.1 I.iU&... 1.1 L.a~ ... -- - - 1I1111.2~ 111111.4 111111.6 111111.25 111111.4 111111.6 MICROCOPY RESOLUTION TEST CHART MICROCOPY RESOLUTION TEST CHART NATIONAL BUR" I OF STANDARDS·1963·A NATIONAL BUREAU OF STANDARDS·1963·A A Taxonomic Revision of the Genus Lolium By EnwAim E. TERRELL Crops Research Division Technical Bulletin No. 1392 Agricultural Research Service UNITED STATES DEPARTMENT OF AGRICULTURE CONTENTS ,PAOB Introduction ------------------------------------------- 1 Cytology and genetics --________________________________ 3 Taxonomic and evolutionary relationships ___________------- 4 Systematic treatment ------_____________________________ 5 Key to mature and complete plants __________________ 6 1. Loli-um pe'renne L. --______________ . _._______________ 7 2. Loli1tm multijlo1"Um Lam. __________________________ 10 3. Lolium rigid1tm Gaud. ____________________________ 15 4. Lolium 8'ltbulatum Vis. ____________________________ 26 5. Loli1tm cana1'iense Steud. _________________________ 30 6. LoUum temulenium L. ____________________________ 35 7. Loli1tm 1'emot'lL'rn Schrank _________________________ 38 8. Lolium pen;iC1tm Boiss. & Hohen. ex Boiss. __________ 41 Literature cited --------________________________________ 44 Appendix ------------__________________________________ 46 Synonyms -----_________________________________ .___ 46 Names under Loli1Lm referring to hybrids ______________ 58 Dubious names ------_______________________________ 59 Excluded names --__________________________________ 59 Index to names ------__________________________________ 60 ., \ Washington, D.C. Issued August 1968 For 9111e by the Superintendent of Documents, U.S. Goverument Printing Office \Vnshington, D.C. 20·102 - Pric~ .10 cents ii A Taxonomic Revision of tbeGenus Lolitun By EDw,um E. TBHlUlLL, ?'csc(t'rch botan'jst Orops Resowrch Division, }l!l/'!Cltltlt'l'al Reseal'eh Service lNTUODUCTION Loa'wn, a genus of the gras~ 'mnllijlora;ln, and L, 'rigicl'lL11L, can (amily, includcs eight species, ac­ be clearly separated from the au­ cording to the present treatment. togamolls group, L. 1.{,IJ1!uleni:n?n The species are indigenous to and L. /'CIIW!lWL. Loliu11l. 'Il/;ult-i­ Europe, the North Atlantic ,[s­ flo)'IWL and L. rigid'WlIL nre like L. lands, temperate Asia, and north /('l/i.ul{')'l! Wit and L. 1'C?lwL'nm in Africa, but they ha\'e been widely being separable from each other distributed to other parts of the only with ditliculty. LoUwn pe­ world. T\\'o members of the genus r('nne cliffers more from L. 'I'igi­ are huporlanL economic grasses clwn than it does from L. multi­ in cool temperate climates flamm. throughout the world. 1n EUrope The chHotic stat.e of the nomen­ L. 1Jc!'('nne (perennial L-yegl'<.lSs) clature of Laliwn conside1'ably and L. 111 uti ijlo/'/o)L (annual rye­ hampered efforts to dedse a bet­ grass, I Lalian ryegTHss) are espe­ ter classification; some order had cially :mportanl as JOl'age grasses. t.o be made of the approximately .Tn the Uniled SL<ltes perennial rye­ ·tHO published names. Many names grass i::; sometimes grown in per­ additional to those listed in "In­ manenL pastures, espcciallyin the dex Kewensis" anel the Smith­ Pac:ific Coast ane! Southern States. sonian "Index to Grass Species" It is a coml~lon but minor compo­ were found during this study; un­ nent of Inwn grass m ixtul'es. In doubLedly there are others undis­ the SO~ILhel'n Vnited StaLes an­ covered, because Loti'n'ln has I)een nual ryegl'ass is sO\\'n in the fall treated in many European floras La furnish a good temporary lawn in the past 200 years. All of the in winLer and spring. It may be known published names are given used also ~lS excellent temporary in an appendix, organized in the pasture or cut as hay. usual chronological sequences to­ . There are no modern taxonomic gether with some nomenclatural revisions of the genus, HOllville and morphological annotations. A (J 853) I in H doctoral thesis recog­ large proportion of the published nized three broad species, Essad names refer to unimportant local (195;1) stuclied fiye species by vaI"iants, environmental modifica­ means of discriminant functions. tions, abnon~al forms, or are He concluded that the allogamolls laLel' homonyms. group, including L. pcrennc, L. The present study represents an "alpha taxonomy" stage in the I Sec Literature Cited, p, ,111. accrual of knowledge concerning 1 2 TECHNICAL BULLETiN 1392, U.S. DEPT. OF AGRICULTURE the genus. Man's cultivation, Although I collec,ted some spec­ along with the weedy nature of imens in the Eastel'll United the genus, has resulted in com­ States and in POl'tugal and Spain plex evolutionary patterns. Infor­ and gl'ew small numbers of plants mation has been obtained by cy­ in the greenhouse, herbarium tological and genetical studies on the genus in cultivation, but very specimens provided most of the little data exist about natural data for this il1veE'ltigation. Abl;mt populations in the l'ileditenanean 4,780 specimens were received on andsouthweste1'n Asian regions loans. -the centers of variation. Field collections obtained and analyzed In addition, I visited certain by modern methods are needed to ElU'opean and American herbaria l'eveal variation pattel'lls. Until to examine types and entire col­ more evidence is available con­ lections. cerning natural hybridization In the following list (abbrevia­ and other phenomena, a more use­ ful classification is achieved by tions by Lanjouw and Stafleu, recognizing the e::-"'1;remes as spe­ 1964) herbaria fl'om which loans cies anel at the same time calling were received are designated with attention to intergradations be­ "(L)" and those visited are desig­ tween them. nated "(V)" : BAG National Herbarium of Iraq, Baghdad (L) BC Instituto Botfmico de Barcelona, Barcelona (V) BM British Museum (Natural History), London (L, V) CAL Central National Herbarinm, Calcutta (L) COl Botanical Institute of Univel'sity of Coimbra, Portugal (L) DAO Department of Agriculture, Ottawa, Ontario (L) E Royal Botanic Garden, Edinburgh (L) FI Istituto Botanico, Florence (L, V) G Conservatoire et Jardin Botaniques, Geneva (L, V) GH Gray Herbarium, Cambridge (V) HUJ Hebrew University, Jerusalem (L) K R(lyal Botanic Gardens, Kew (L, V) LAG Musee Botanique Cantonal, Lausanne (L) LE Komarov Botanicallnstitute, Leningrad (L) LINN Lil1nean Society, London (V) LISC Centro de Bohlnica da Junta de Investigag6es do Ultramar, Lisbon (L, V) LISE Esta~ao Agronomica Naciona1, Oeiras, Portugal (V) LIS! Instituto Superior de Agronomia, Lisbon (V) LISU UniYersity of Lisbon, Lisbon (V) MA Instituto "Antonio Jose Cavanilles", Madrid (V) l\{ADIVI Museu IVlunicipal do Funclml, :Madeil'a (L) MADS Museu de Historia Natural do Seminario do Funchal, Madeira (L) NA U.S. National Arboretum, Washington (L, V) NY New York Botanical Garden, New York (L, V) ORT J al'c1iu de Aclimatacion de 1a Orotava, Tenerife, Canary Islands (L) A TAXONOMIC REVISION OF THE GENUS LOLIUM 3 P Museum. d'Histo~1!e Natur~l1e, Paris (V) PR National Museum, Prague (L) RAW Gordon College, Rawalpindi, Pakistan (L) US U.S. National Herbarium, Washington (L, V) W Naturhistorisches Museum, Vienna (L) CYTOLOGY AND GENETICS Artificial crosses made by ge­ hybrids of L. po'renne X F.pm­ neticiets provide some information tens'is are particularly frequent in about the biosystematics of Lo­ Europe. liunl.. 'rhe general facts from this According to geneticists, L. 'rig'i­ source were reviewed (Terrell d'win val'. ?"ig'idum, is self-incom­ 1966) and may be briefly sum­ patible whereas L. ?"ig'id~tm val'. marized as follows: (1) All taxa '1'ottuollio'ides is self-compatible. for which chromosome numbers This inconsistency may be more have been counted are diploid apparent than real because of in­ \vjth 2nc.::1Ll; (2) L. pel'enne, L. sufficient samples studied as well 'nmltijloYllln, and perhaps L. ?'ig'i­ as uncertainties regarding taxo­ cl'U/)i are self-incompatible and nomic identities. cross~pollinated; (3) L. temulen­ The ability of certain Lolium tUlIl, L. nmwhun, and perhaps L. species to cross with FestucCL sec­ rigichun val'. 'I'ottuollio';cles (L. tion Bovinae suggests to certain loliacclLln) are self-compatible authors that the genera should be and self-pollinated; (4) all taxa united (Terrell 1966). Their mor­ of Lotium are more Ol' less inter­ phological differences (spicate vs. fertile; (5) most of the species of paniculate; one glume vs. two Lolium are known to cross with glumes) may be less significant Fcsflwfl section Bovinae, including than heretofore believed. However, F. PI'(l {cnsis Huds. (F. cZCLtio?' L.) , the species of Lolium are more F. (L'/'nnclinacea Sehreb., and F. closely related to each other than giyantca (L.) ViII.; (6) certain to Fest'llGCL. I favor, at least in the species of Loliun~ cross with con­ present state of knowledge, the siderable difliculty with FestucCL retention of LoUnm and Festuca section Festllca, including F. ovinCL as two distinct genera. That they L., F.mul'Cl L., and their close do cross in nature as well as arti­ relatives; (7) Loliwn has never ficially does not necessarily mean been crossed successful1y with any they are congeneric. Similar situ­ other genus besides Fcsh~ca; (8) ations are known in other grass Lolium and Frstuca constitute a genera. Each f!ase must be decided comparium because they can c.'(­ on its own merits. More facts are change genes; and (9) in contrast needed about hybridization and to LoUllIn, Fesluc(l includes poly­ morphology to arrive at general ploid taxa. principles regarding generic lim­ Natural hybl'ids of various Lo­ .its in grasses. Classification should Ullin species and intergenel'ic hy­ also provide a stable and usable brids with Fe.'l/'uc(( have been nomenclature. Name changes found, chiefly in Europe (briefly should be made only when the evi­ listed by Terrell 1966).