N. MANCHESTER, INDIANA w. sv aS VOLUME 64 ANNALS OF THE MISSOURI BOTANICAL CARDEN Published by the Missouri Botanical Garden Press, St. Louis, Missouri 63110. © Missouri Botanical Garden 1978 CONTENTS Apams, Rosert P. Chemosystematics—Analysis of Populational Differen- tiation and Variability of Ancestral and Recent Populations of Junip- erus ashei AsHTON, P. S. A Contribution of Rain Forest Research to Evolutionary Theory AusriN, DANIEL. F. Realignment of the Species Placed in Exogonium ( Convolvulaceae ) AVERETT, JOHN E. Taxonomic Notes and New Combinations in Leucophy- salis (Solanaceae) AVERETT, Jonn E. Chemosystematics: The Twenty-third Systematics Sym- posium Broome, C. Rose. (see Stibolt, Virginia M., C. Rose Broome & James L. Reveal) CARLQUIST, SHERWIN. Wood Anatomy of Onagraceae: Additional Species and Concepts D'Arcy, WILLIAM G. (see Gentry, Johnnie L., Jr. & William G. D'Arcy) DavipsE, Gerrit. New Taxa and Combinations in the Genus Lasiacis Gà»)t % e . S c iai DavipsE, Gerrit. (see Goldblatt, Peter & Gerrit Davidse) |... Dietrich, WERNER. The South American Species of Oenothera Sect. Oe- nothera (Raimannia, Renneria; Onagraceae) : Dunn, Davin B. & WILLIAM E. Harmon. The Lupinus montanus Complex of Mexico and Central America Eype, RICHARD H. Reproductive Structures and Evolution in Ludwigia Onagraceae. I. Androecium, Placentation, Merism FAIRBROTHERS, Davin E. Perspectives in Plant Serotaxonomy FEENv, PAUL. Defensive Ecology of the Cruciferae Gentry, ALwyn H. A New Jacaranda (Bignoniaceae) from Ecuador and Peru. Gentry, ALwyn H. A New Species of Bignoniaceae from Madagascar o Atwyn H. New Records of Apocynaceae for Panama and the Chocó Gentry, ALwyn H. New Species of Gibsoniothamnus (Scrophulariaceae/ Bignoniaceae) and Tournefortia (Boraginaceae) from Eastern Panama and the Choc Gentry, ALWYN H. Studies in Bignoniaceae 25: New Species and Com- binations in South American Bignoniaceae Gentry, JOHNNIE L., Ja. & WILLIAM G. D'Arcy. Solanum armentalis: A New Species from Costa Rica 311 376 GorpBLATT, Perer. Chromosome Number in Pillansia (Iridaceae) — GorpBLATT, PETER. Herbertia (Iridaceae) Reinstated as a Valid Generic Name GorpBLATT, PETER. Systematics of Moraea (Iridaceae) in Tropical Africa GorpBLATT, PETER & Gerrit Davipse. Chromosome Numbers in Legumes Harmon, WILLIAM E. (see Dunn, David B. & William E. Harmon) - Hocu, PETER H. & Perer H. Raven. New Combinations in Epilobium (Onagraceae) UU U UU au S pn DT Huckins, CHARLES ALBERT. In Chromosome Numbers of Phanerogams. 7 Janzen, DANIEL H. Promising Directions of Study in Tropical Animal- Plant ne oe tome roc MM í Jorpan, Cart F. & ERNESTO MEDINA. Ecosystem Research in the Tropics Kinc, Many-CLamr. The Applications of Molecular Evolution to System- atics: Rates, Regulation, and the Role of Natural Selection KING, ROBERT M. & Hanorp RoBINsoN. Guayania davidsei and Hebecli- nium gentryi, New Species from Northern South America (Eupato- eee) 8 Manny, Tow J. The Order Centrospenmae —-— — o c e MaciLL, ROBERT E. (see Seavey, S. R., Robert E. Magill & Peter H. Raven) MEDpINA, Ernesto. (see Jordan, Carl F. & Ernesto Medina Gorrum, L. D. Electrophoretic Evidence and Plant Systematics —_ PARNELL, Dennis R. (see Raven, Peter H. & Dennis R. Parnell) PRANCE, GHILLEAN T. Plant Inventory of the Tropics: Where Do We Stand? .. FU Ramirez B., WILLIAM. A New Classification of Ficus Raven, Perer H. A New Species of Lopezia (Onagraceae) from Sinaloa, Mexico DINE Raven, Peter H. Perspectives in Tropical Botany: Concluding Remarks Raven, PETER H. (see Hoch, Peter H. & Peter H. Raven Raven, PETER H. (see Seavey, S. R., Robert E. Magill & Peter H. Raven) RAvEN, Perer H. (see Tomlinson, P. B. & Peter H. Raven) Raven, Peter H. & Dennis R. PARNELL. Reinterpretation of the Type of Godetia bottae Spach (Onagraceae) REVEAL, JAMES L. (see Stibolt, Virginia M., C. Rose Broome & James L. Reveal) Rosinson, HARoLp. (see King, Robert M. & Harold Robinson Rourke, JOHN & DELBERT Wiens. Convergent Evolution in South African and Australian Proteaceae and its Possible Bearing on Pollination by Nonflying Mammals Seavey, S. R., ROBERT E. MAGrLL & PETER H. Raven. Evolution of Seed Size, Shape, ad toes poor rim in the Tribe Epilobieae (Ona- graceae) SPELLMAN, Davin L. Four Species of Asclepiadaceae New to Panama STIBOLT, VIRGINIA M., C. ROSE Broome & James L. A Alnus mari- tima Muhl ex Nutt., not Alnus metoporina Furlo STRALEY, GERALD B. Systematics of Oenothera Sect. = (Onagraceae) ToMuinson, P. B. Plant Morphology and Anatomy in the Tropics—The Need for Integrated: —— mt A TEENS ToMrLINSON, P. B. & PETER H. RAvEN. Perspectives in Tropical Botany: Introduction Towner, Howard F. The Biosystematics of Calylophus (Onagraceae) .. Turner, B. L. Chemosystematics and its Effect upon the Traditionalist . Wiens, DELBERT. (see Rourke, John & Delbert Wiens) WITHERSPOON, JouN T. New Taxa and Combinations in Eragrostis (Poa- ceae) WUNDERLIN, RICHARD P. A New Species of Bauhinia (Leguminosae) from Peru 371 Mob eb gs n Ress 55 VER 8 . “Sasa VOLUME ó4 1977 NUMBER 1 The ANNALS contains papers, primarily in s botan. = contributed from the Missouri Botanical Garden. Papers originating outside the Garden will also be accepted. Authors should write the editor for inf i 8 for i in = the ANNALS. E: EprrontaL C EC Giner Dave. Editor-i chef X Missouri Botanical Garden W. G. D'Arcy, Editor Flora of Panama Missouri Botanical Garden JohN D. DwYER Missouri Botanical Garden c St. Loud University | PETER GOLDBLATT Missouri Botanical Garden Published four times a year by the Missouri Botanical Garden Press, St. Ae pitis Missouri 63110. For subscription information contact the Business Office of the Annals, P.O. Box 368, 1041 New Hampshire, Lawrence, Kansas 66044 Subscription price is $40 per volume U.S., Canada, and Mexico, $45 all other countries. Four issues per volum Second class postage paid at Lawrence, Kansas 66044 O Missouri Botanical Garden 1977 ANNALS OF THE MISSOURI BOTANICAL GARDEN VOLUME 64 1977 NUMBER 1 CONVERGENT FLORAL EVOLUTION IN SOUTH AFRICAN AND AUSTRALIAN PROTEACEAE AND ITS POSSIBLE BEARING ON POLLINATION BY NONFLYING MAMMALS Joun Rourke! AND DELBERT WIENS” ABSTRACT Striking convergent evolution for a hidden (cryptic), ground flowering (geoflorous) habit in distantly related, low shrubby Australian and South African Proteaceae is interpreted as an adaptation for pollination by nonflying mammals. The cryptic, geoflorous habit is age cially well de veloped in species groups of Dryandra in southwestern Australia and Protea n the Cape region of South Africa. Considerable circumstantial evidence exists in both re- um for pollination by mouselike, often arboreal marsupials in Dryandra and true rodents in Protea. Evidence from inflorescence structure suggests the cryptic, 5 habit is de- rived from bird-pollinated species, possibly in response to fires common in the sclerophyllous communities where these genera grow. A number of floral char: ae and the occurrence in Australia of mouselike marsupials adapted to a nectar (and pollen?) diet suggests that a n of flowers has evolved for po ined by nonflying mammals. This postulated floral class sibly also extends to other Australian arb n jroteaceous and also myrtaceous genera, Ent in South Africa is probably restricted to Protea Pollination by nonflying mammals is largely ignored or given little credence in current treatments of pollination ecology ( Faegri & van der Pijl, 1971; Proctor & Yeo, 1972). There is, however, good reason for this; all the available evidence relating to this phenomenon is either circumstantial, inferential, or anecdotal. Nonetheless, field observations in Australia and South Africa and a subse- quent search of the literature have led us to believe that true rodents and mar- supials may, in fact, be the normal pollinators of several southern hemisphere proteaceous genera. Furthermore, various floral characteristics in these genera and the special adaptations for nectar feeding in some of the putative pollinators suggest structural coadaptations by both flowers and apparent pollinators. Al- though plans are underway to conduct definitive studies, no unequivocal evidence can be presented at this time for regular pollination by nonflying mammals, and ! Kirstenbosch Botanic Garden, Newlands, Cape Province, South Africa 7700. ? Department of Biology, University of Utah, Salt Lake City, Utah 84112, U.S.A. ANN. Missounr Bor. Garp. 64: 1-17. 1977. 9 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 that is not the intent of this paper. We hope, however, that our comments will help to reopen and stimulate research in this fascinating area of pollination bi- ology pioneered by Porsch (1934, 1935, 1936a, 1936b) and subsequently neglected for 40 years. The purpose of this paper is fourfold: (1) to elucidate our observa- tions and ideas on inferred pollination by nonflying mammals in the South African and Australian Proteaceae, (2) to point out the striking convergent evolution of flowering habits between southwestern Australian and South African Proteaceae of the Cape region, (3) to review some of the rather scattered and fragmentary literature on the subject, and (4) to evaluate the evidence for the existence of a class of flowers adapted to pollination by nonflying mammals. References to the subject of pollination by nonflying mammals usually men- tion the arboreal Australian marsupials which apparently feed on nectar (e.g. the honey possum, Tarsipes spencerae) and to introduced rats suspected of pol- linating a climbing pandan ( Freycinetia arborea Gaudich.) in Hawaii. Faegri & van der Pijl (1971) and Proctor & Yeo (1972) furthermore state that no flowers appear to be adapted for pollination by nonflying mammals, although Faegri and van der Pijl mention the classic papers by Porsch (1934, 1935, 1936a, 1936b) in which he builds a case for floral adaptations to pollination by nonflying mam- mals in several Australian genera. Grant (1950) mentions, without comment, a "marsupial" pollinated flower class based on the proteaceous genus Dryandra. In addition to rodents and mouselike marsupials, some primates may also be regular pollinators. For example, according to Coe & Isaac (1965) the baobab (Adansonia digitata L.) is pollinated in East Africa by the lesser bush baby (a lorisid primate). This characteristic African tree is generally considered to be bat pollinated. Petter (1962) mentions that several arboreal, mouselike lemurs ( Lemur, Varecia, Hapalemur, Microcebus) visit flowers seeking nectar and are generally attracted by sweet liquids in captivity. More recently Sussman & Tat- tersall (1976, and personal communication) demonstrate that Lemur mongoz mongoz is apparently an important pollinator of introduced kapok (Ceiba pen- tandra Gaertn.) in Madagascar. F. L. Carpenter (personal communication) has data from Australia indicating that some species of Banksia are pollinated almost entirely by nonflying mammals, including an indigenous rat (Rattus fuscipes) and various marsupials. It is not our intent to evaluate the entire literature here. There are, however, numerous instances of various mammals being observed on or around flowers ( Porsch, 1934), but the nature of their activities are, in fact, virtually unknown. As Faegri & van der Pijl (1971) point out with respect to pollination by non- flying mammals *much research remains to be done to establish ic be- tween possible regular pollinators and the blossoms in which they wor FLORAL CHARACTERISTICS AND CONVERGENT EVOLUTION OF PROTEACEAE PUTATIVELY POLLINATED BY NONFLYING MAMMALS The most obvious Proteaceae are trees and large shrubs, e.g., Grevillea and Banksia in Australia, and Protea in South Africa. Less known, however, is the occurrence of species groups on both these continents with inflorescences at or near ground level (geoflorous) and typically obscured from external view by 1977] ROURKE & WIENS—NONFLYING MAMMAL POLLINATION 3 overlying foliage (cryptic). The taxonomic distribution of these cryptic, geo- florous species is limited principally to two distinct sections of Protea [Hypo- cephalae and Microgeantheae, sensu Phillips (1912)] and some additional species of uncertain sectional classification in the Cape region of South Africa; in south- western Australia, however, this flowering habit is associated with at least five genera (Banksia, Conospermum, Dryandra, Isopogon, and Petrophile) but is best developed in Dryandra [series Aphragmia and Niveae, sensu Bentham (1870) and to a somewhat lesser extent in Banksia. In these equivalent infrageneric groupings in Protea and Dryandra the growth habit is low, tufted, and often rhizomatous. The flowers occur in heads, usually at ground level, or occasionally up to 30 cm high, but in either case the heads are typically deeply hidden within the foliage of the dense and widely spreading branch systems. The heads are generally visible only if the branches are forcibly parted and the base of the plant carefully examined (Figs. 1-6). The flowers are surrounded by a prominent series of overlapping bracts forming a cup-shaped involucre. The bracts vary in color through various shades of brown and are often flushed with different dull reddish tints. An inflorescence contains per- haps 100-200 flowers, but the large spikes of Banksia bear several thousand in- dividual flowers. Many of the species produce copious amounts of nectar and the heads often emit a distinctive, “nutty” or “yeasty” odor. In the cryptic, geo- florous Cape species of Protea the basal portions of the bracts and flowers, par- ticularly the styles, are also markedly succulent. Excellent illustrations of Protea flowers (but not necessarily the growth habits) can be seen in Rousseau (1970) for South African proteas and in Erickson et al. (1973) for Australian genera. Dryandra, as in most western Australian Proteaceae, develops no obvious suc- culence in the inflorescence or flowers. In general, the geoflorous habit, the cryp- tic positioning of the inflorescences, and the gross (though superficial) morpho- logical similarities of the heads suggest strong convergent evolutionary tendencies. In fact, from a distance one would be hard pressed to distinguish between some species of Dryandra and Protea even though these genera represent the end points of evolution in two subfamilies of the Proteaceae, Grevilleoideae and Proteoideae, respectively, and occur on widely separated continents (Figs. 1-6). EVIDENCE FOR RODENT POLLINATION IN SOUTH AFRICAN PROTEAS Field observations over a period of years of the cryptic, geoflorous species of Protea in the Cape area show that considerable rodent activity is associated with these species (Table 1), but is especially obvious in P. subulifolia. The specific rodent activities associated with this species are: (1) freshly chewed involucral bracts and styles during and just prior to anthesis ( Fig. 7); (2) clearly demarcated networks of heavily used runways linking different plants within populations, and which often intertwine around flowering and old fruiting heads; and (3) occa- sional burrows at the base of the plants. The runways and burrows are related to activities of the Cape striped field mouse (Rhabdomys pumilio pumilio). On various occasions and in different populations this animal (which is diurnal) was observed on runways between 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 5 y — eid 1977] ROURKE & WIENS- NONFLYING MAMMAL POLLINATION 5 TABLE I. The South African species of Protea in sections Hypocephalae and Micro- geantheae. All species Bode flowering or near ground flowering. H lypoc lun: ^M "mayqan wass P. st ra (Salish. ex P. acaulos (L.) Reichard’ P. montana E. Mey. ex Meisn. Knight) Rourke* P. angustata R. Br. . restionifolia (Salisb. ex P. amplexicaulis R. Br. P. aspera Phillips? Knight) Rycroft’ P. decurrens Phillips P. cordata Thunb. P. revoluta Buek ex Meisn. P. humiflora Andrews’ P. foliosa Rourke P. scabra R. Br. P. glaucophylla Salisb. P. scabriuscula Phillips P. intonsa Ro d P. scolpendrium R. P. laevis Thun F. scorzone rifolia Salish. P. lorea R. Br. gh P. 0 Phillips“ P: vogtsiae Rourke* a Species in which evidence of rodent activities has been observed on flowers b Inclusion in this taxonomic section questionable flowering plants of P. subulifolia. That the Cape striped field mouse is attracted to the cryptic, geoflorous inflorescences of P. subulifolia was further demon- strated when it was live-trapped utilizing fresh flowering heads of this species as bait. In this instance traditional rodent baits such as peanut butter were in- effectual in capturing this animal. The Cape striped field mouse apparently also visits the flowering heads of Protea nana (Berg.) Thunb., a low shrubby species with pendulous heads of un- certain pollination type and not a member of the geoflorous sections. The soft floral parts of P. nana were chewed in the same manner as P. subulifolia and a Cape striped field mouse was trapped at this plant within 24 h of the first noted rodent activity e fleshy involucral bracts and styles of the cryptic, geoflorous proteas show widespread evidence of being chewed. In one population of P. subulifolia 17 plants bearing 49 inflorescences with open flowers were observed in an area of approximately 30 m?; 20 heads, or 40%, showed extensive evidence of chewed bracts and styles ( Cape striped field mice were common in the area). The con- sistent occurrence of chewed bracts and styles in the cryptic, geoflorous proteas suggests that they function as food bodies; supporting this idea is the sweetness (at least to the human palate) of these structures. In the large, bird-pollinated proteas the bracts not only lack succulence but are markedly acrid, apparently containing high concentrations of tannin. In spite of their sweet, fleshy nature and apparent lack of tannin, all the bracts and mature styles of any head are rarely eaten. That the inflorescences are not completely destroyed suggests the presence of secondary compounds which might limit the amount of feeding as proposed by Freeland & Janzen (1974 < —]. General aspect 2 1 p just beginning to flower (near Hermanus, d Prov., South Africa).—2. Cr , geoflorous inflorescences of P. MN (same plant as shave); —3. General aa of Dr andra tenuifolia in full flo (Stirling Range, West Australia).—4. Cryptic, geoflorous ences of D. ter ola e plant as above). 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Sweet, fleshy bracts are also associated with pollination in Freycinetia ( Pan- danaceae) where they reportedly attract introduced rats in Hawaii (Degener, 1945) and in other areas of the Pacific (B. C. Stone, personal communication ). Freycinetia insignis Blume, an Asian bat-pollinated species, apparently utilizes only odor and fleshy bracts as attracting devices (Proctor & Yeo, 1972 The sense of smell is well developed in rodents, and in view of the hidden nature of the inflorescences, odor must be the primary attracting mechanism re- gardless of the pollinator. Furthermore, at the time of flowering in P. subulifolia (late winter), the fleshy bracts and styles constitute one of the best sources of soft palatable vegetable matter in the local plant community. Thus flowering might correlate with the low point in the food cycle of rodents. The Cape striped field mouse is apparently fond of soft vegetable matter, sometimes becoming a nuisance in vegetable gardens (Roberts, 1951). As further evidence of this dietary habit, newly harvested shoots of another proteaceous shrub, Leucodendron modestum Williams, were observed along a typical runway and entrance to a Cape striped field mouse burrow. If the chewing activity in the flowering heads of the Cape species of geoflorous proteas is due to the Cape striped field mouse, or a similar-sized animal, pollen would surely accumulate about the head of the animal and should theoretically be capable of transfer to nearby plants. The fur of mammals should provide an excellent surface for pollen accumulation. This is demonstrated by the presence of pollen on the head of a nectar-feeding Aus- tralian marsupial, the sugar glider (Petaurus breviceps) (Breeden & Breeden, inside back cover). An interesting description of pollen accumulation on the Australian honey possum (Tarsipes spencerae) feeding on Proteaceae is also given by Vose (1972). Because of its known association with Protea (especially P. subulifolia), the Cape striped field mouse is perhaps the best possibility for a mammal pollinator of the cryptic, geoflorous species of Protea. However, other rodents in the Cape fauna should also be examined for possible activities relating to pollination. Dr. J. Jarvis of Cape Town University (personal communication) suggests especially the following animals: Dendromus melanotis (climbing mouse), Leggada minu- toides (dwarf mouse), Otomys irroratus (vlei otomys), and Acomys subspinosus ( Cape spiny mouse). None of these animals, however, appear to have any special adaptations for nectar or pollen feeding. A single case of interspecific hybridization (P. restionifolia x P. humiflora) is known among the cryptic, geoflorous species of Cape Protea. That such a cross occurs is proof that pollen can be transferred between these species. Further- more, evidence of rodent activity is known in both parental species of the cross. The Proteaceae are apparently adapted for outcrossing and thus require a mecha- nism for pollen transfer. The family is apparently either protandrous (Rao, 1971) or self-incompatible (Horn, 1962). Pollen dispersal ultimately occurs from a specialized region of the style apex known as the pollen presenter ( Rourke, 1969). This is so close to the slitlike stigmatic surface that mechanisms to prevent autogamy must be present or selfing would be the rule and pollination unneces- sary. 1977] ROURKE & WIENS—NONFLYING MAMMAL POLLINATION 7 EVIDENCE FOR MARSUPIAL AND RODENT POLLINATION IN AUSTRALIAN PROTEACEAE Field observations of the inflorescences of Dryandra tenuifolia R. Br. in south- western Australia also showed evidence of mammal activities similar to those men- tioned for Protea subulifolia from the Cape region of South Africa; chewed heads were particularly common. The inflorescences were also odoriferous and the scent was surprisingly similar to the “yeastlike” odors prevalent in the cryptic, geoflorous species of African Protea. Copious nectar was not detected, but our observations were made in mid-afternoon when nectar content was possibly low. Nectar production in the Australian cryptic, geoflorous Proteaceae may be largely nocturnal to coincide with increased animal activity at that time (Mor- combe, 1968). Porsch (1935) repeatedly mentions high nectar production in Dryandra nivea R. Br., which he observed under cultivation. He also noted noc- turnal anthesis and an odor of “sour milk” or “caraway liquor" in this species. Dr. Alex George (personal communication) has also seen apparent mammal activity in the inflorescences of the cryptic, geoflorous species of Banksia where chewing and disturbance of the flowers appeared to be similar to our observations in South Africa. F. L. Carpenter (personal communication) also has interesting evidence that Banksia species in eastern Australia are largely pollinated by nonflying mam- mals. She correlates nonflying mammal pollination in Banksia with the occur- rence of stiff inflexed styles (illustrated in Baglin et al., 1972) which apparently exclude foraging birds. Porsch (1935) suggested this as a feature of marsupial- pollinated banksias; he also proposed that the “basket”-like inflorescences in some dryandras were adapted to accommodate the heads of various marsupials ( Fig. 8). The situation in Australia, however, is probably more complex than in South Africa. For example, many of the large, shrubby and even arboreal Proteaceae (and also Myrtaceae) are also visited by nonflying mammals in addition to the cryptic, geoflorous species, yet the latter appear to be better adapted for polli- nation by nonflying mammals. Most workers probably consider these nonground flowering species to be bird pollinated (e.g., Carlquist, 1974). Admittedly many of the floral characteristics of genera such as Banksia do suggest bird pollination. Yet some traits clearly do not. For example, Baglin et al. (1972) state that all Banksia inflorescences are odoriferous, yet odor is not associated with ornithophily. Additionally, Morcombe (1968) reports that in Banksia nectar secretion is pro- lific at night, a condition hardly adapted to pollination by diurnal flower birds. Morcombe suggests that the great abundance of nocturnal insects are attracted to Banksia inflorescences by the copious nectar, and these in turn are what entices nonflying mammals to the flowers. Considering the highly specialized adapta- tions of an animal such as the honey possum (see following discussion) for a nec- tar (and pollen?) diet, it seems unlikely that insects would be the prime at- tractant, at least for this animal. However, animals such as the southwestern bush rat (a true rodent) might well be attracted by insects. But this would hardly explain why nectar secretion is abundant at night, since insects are highly unlikely pollinators of these flowers. Typically, nectar secretion is synchronized temporally for visitation by the established pollinators coadapted to that par- ticular flower ( Faegri & van der Pijl, 1971). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 1977] ROURKE & WIENS—-NONFLYING MAMMAL POLLINATION 9 DERIVATION OF CRYPTIC, GEOFLOROUS SPECIES FROM ORNITHOPHILOUS PROTOTYPES Various sunbirds and the Cape sugarbird are the typical pollinators of the well-known shrubby proteas in the Cape region with large terminal inflorescences (Fig. 9). However, the cryptic, geoflorous positioning of the inflorescences in sections Hypocephalae and Microgeantheae must preclude bird pollination, since birds are attracted to flowers visually (Raven, 1972) and odor is not a characteristic of bird-pollinated flowers. In fact, sunbird or sugarbird visits to the cryptic, geoflorous proteas would violate established behavioral patterns in these birds. They are not generally known to frequent the ground, or to explore the dense interior of low shrubs which do not have exposed, colorful flowers. Further evidence that birds are unlikely pollinators of the cryptic, geoflorous proteas is based on observations of P. nana. As previously mentioned, this species has pendulous, relatively small (ca. 3 cm wide), dark reddish heads. Initially one might assume that they were bird pollinated. However, a number of flower- ing plants of P. nana were observed in an area densely populated by the orange- breasted sunbird, Nectarinia violacea and the Cape sugarbird ( Promerops cafer) which were feeding freely on several proteaceous shrubs with large terminal heads, and some ericas; however, no birds were observed on P. nana. Because the heads are pendulous, flower-visiting birds would probably have to hover to ob- tain nectar. Sunbirds are capable of hovering (Skead, 1967) but unlike hum- mingbirds, they hover clumsily. Normally they feed while clasping branches. Nonetheless, in an area with a high density of nectar-feeding birds, a great va- riety of flowers are normally visited in addition to the preferred species. If sun- birds and sugarbirds showed interest in the flowers of P. nana, at least rare visits to these plants would be expected. If nectar-seeking birds are not attracted to P. nana, whose inflorescences are visually conspicuous but otherwise generally re- semble the cryptic, geoflorous species, it is still more difficult to believe that birds pollinate the latter group. In fact, the great majority of these South African Proteaceae with dark reddish bracts and mostly pendulous flowers might well be pollinated by nonflying mammals. The cryptic, geoflorous proteas do retain the copious nectar supply typical of bird flowers. They differ from bird flowers, however, by (1) bearing their flowers at or near ground level in a hidden position, (2) emitting a strong “yeast- like” odor, (3) possessing much shorter flowers (ca. 1.5 cm high), and heads of smaller diameter (ca. cm wide), and (4) the dull purplish brown coloration of the heads as opposed to the bright, vivid red and/or yellow inflorescences of the bird-pollinated species. Essentially the same arguments apply to the situation in Australia, except 1 5-8.—5. Inflorescences of Protea subulifolia at anthesis (same plant as Figs. 1-2).—6. In 5 of Dryandra tenuifolia at an DER (same plant as Figs. 3-4).—7. P. subulifolia showing chewed 1 bracts and styles (left) and intact inflorescence (right) rcd e Vlei, — Prov., Sout rica š fatica sp., note inflexed styles forming 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 1977] ROURKE & WIENS—-NONFLYING MAMMAL POLLINATION 11 that here the predominant flower birds are the honey eaters (Meliphagidae). Likewise in many of the critical genera such as Banksia, some species are ap- parently adapted for bird pollination and others for pollination by nonflying mammals. Although we believe the cryptic, geoflorous Proteaceae are not pollinated by birds, the system nonetheless appears to have evolved from a bird-pollinated prototype. In addition to some general aspects of the inflorescence and the copious supply of nectar, the nature of the branching patterns associated with inflorescence development also supports the derived nature of the cryptic, geo- florous Proteaceae. The large, shrubby bird-pollinated proteas have terminal inflorescences, whereas the inflorescences in the geoflorous species are largely axillary. These axillary inflorescences are almost certainly derived from terminal inflorescences by progressive stem reduction. Members of the section Pinifoliae (which includes P. nana) possess intermediate forms in which the stem bearing the inflorescence is greatly shortened. Complete reduction of this stem would give rise to the almost sessile, apparent axillary inflorescences characteristic of P. subulifolia and other highly reduced types that occur in the geoflorous sections. Thus morphological evidence supports the proposition that the geoflorous species are derived types originating from bird-pollinated groups. Both L. A. S. Johnson and A. George (personal communications) support the notion that the cryptic, geoflorous Australian Proteaceae are also derived types. Another interesting example of the apparently derived nature of the cryptic habit occurs in Protea recondita Buek ex Meisn. where floral crypsis is accom- plished through an entirely different mechanism than geoflory. This species is a low shrub (up to perhaps 1 m high) with terminal inflorescences positioned similarly to the bird-pollinated proteas. The bases of the heads, however, are encircled by a cluster of unusually large, vertically oriented leaves (bracts). These bracts enfold the entire inflorescence (rather like a cabbage!) and, in effect, obscure the head from external view during anthesis (Figs. If the cryptic, geoflorous species of Proteaceae are adapted for pollination by nonflying mammals and were derived from bird-pollinated prototypes, what selective forces might have shifted the system in this direction? The ecological community in which these plants occur provides a possible explanation. Both the Cape region and southwestern Australia are essentially sclerophyllous, fire- adapted shrub communities. In fact, the general aspect of the two communities is remarkably similar, even to the characteristic brownish cast of the vegetation. Furthermore, both regions are extraordinarily rich floristically. Only tropical rainforests are apparently richer in plant species diversity. Both floras are also < Ficures 9-12.—9. Protea compacta, a typical Hee pollinated species at anthesis (near Papies Vlei, Cape Prov., South Africa ).—10. recondita, shoot p hidden terminal in- florescence at anthesis 5 Botanic Garden, Cape Prov. uth Africa).—11. P. re- condita shoot with terminal inflorescence exposed ehind the large, du bracts (same plant as Fig. 10).—12. (left) Banksia ph: (from a kodachrome by Alex George, West Australia ) i cence at anthesis, each rounded point represents one flower, total number of flowers estimated at 4,400. (right) Banksia inflorescence with 34 mature fruits. 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE 2. Australian mammals known to visit flowers to obtain nectar and/or pollen. Kapal. (all marsupials, except Rattus fuscipes) References Acrobates pygmaeus (pigmy lider e & Breeden (1972), Carlquist (1965) Antec hinus apicalis (dibbler Morcombe (1968 A. flavipes (yellow-footed antechinus ) Breeden & Breeden (1972 Duan parvus (mountain pigmy pos- P. Cook ( personal communication ) sum Cercartetus concinnus (southwestern Vose (personal communication), Ride (1970) igmy possum) C. nanus (eastern pigmy possum ) Baglin et al. (1972), Breeden & Breeden (1972) Petaurus australis (fluffy or yellow-bel- Breeden & Breeden (1972) ied glider) P. breviceps (sugar glider) Breeden & Breeden rode 1972), Sleumer (1955) P. norfolcensis (squirrel glider) Breeden & Breeden (1972) Phase vogale tapvatafa (tuan or Wamben- Breeden & Breeden ( Tn ger 1 fuscipes (southern bush-rat ) Morcombe (1968 Tarsipes spencerae (honey mouse) Morcombe (1968), Glauert (1958), Vose (1971, 1972), Ride (1970). characterized by nutritionally depauperate soils (Wild, 1968; Loveless, 1961). Insofar as the evolution of the cryptic, geoflorous habit is concerned, however, we believe the fire-adapted nature of these communities is most important. One way plants can survive burning is to develop rhizomaty. This condition should strongly promote ground flowering. Many, but not all, of the cryptic, geoflorous Proteaceae are rhizomatous. Hence, if rhizomaty has survival value and if there were pollinator competition between bird and nonflying mammals for floral resources, fire and the concomitant development of the rhizomatous habit could have shifted the selective advantage toward nonflying mammal pol- lination. It is also possible, however, that nonflying mammals, as a result of their generally more aggressive behavior, may have simply out-competed birds as pollinators and hence shifted the selective balance in this way. A FLORAL CLASS ADAPTED FOR POLLINATION BY NONFLYING MAMMALS AND EVIDENCE FOR COEVOLUTION Although experimental data are lacking, we believe sufficient circumstantial evidence is available to identify a class of flowers in Proteaceae adapted for pol- lination by nonflying mammals. Furthermore, this class of flowers has evolved independently at least twice (Africa and Australia) and at least one animal, the so-called honey possum (Tarsipes spencerae) has probably coevolved with this floral class in Australia. Furthermore, we believe other animals, particularly some of those in Table 2, may also have coevolved with these proteaceous plants. The fundamental characteristics we believe might distinguish this floral class include: (1) Inflorescences as the basic units of attraction; generally they are cup-shaped heads (spikes in Banksia). (2) Heads typically hidden deep within the foliage, often at or near ground level; if exposed (as in Banksia) then with (a) structural modifications, such as stiff incurved styles or (b) nocturnal 1977] ROURKE & WIENS—NONFLYING MAMMAL POLLINATION 13 rhythms of nectar production and/or anthesis to preclude successful nectar forag- ing by birds. (3) Heads about 2-8 cm wide with perhaps 100-200 flowers (several thousand in Banksia), and strongly attached to stems. (4) Heads producing a copious nectar supply; in some proteas also possessing apparent food bodies in the form of soft, fleshy bracts and styles acting as complementary attractants. (5) Heads odoriferous; we characterize these as “nutty” or “yeasty” in Protea; Porsch (1935) suggests “sour milk” and “caraway liquor” for Dryandra. (6) Heads with reddish brown to purplish bracts, individual flowers mostly whitish. (7) Temporal spacing of anthesis in the inflorescence, thereby limiting the number of simul- taneously open flowers in the head to no more than several of the outer whorls. ost of these characteristics were discussed previously and need no further elaboration. The most obvious feature of this putative floral class is its basic resemblance to bat-pollinated flowers (cf. Faegri & van der Pijl, 1971). The pri- mary differences are the cryptic, geoflorous habits and the compound inflorescence as the attracting unit. There are apparently also structural modifications of the styles in the Australian species to discourage bird foraging. The basic similari- ties to bat flowers, however, should not be surprising since the apparent pol- linators are all small mammals with perhaps generally similar energetic require- ments and sensory systems [Faegri & van der Pijl (1971) point out that echo location is only poorly developed in the flower-feeding bats, Megachiroptera]. One of the strongest lines of indirect evidence supporting the idea of a class of flowers pollinated by nonflying mammals in South Africa and Australia re- volves about the convergent nature of the floral characteristics in these two sub- families of Proteaceae. If one examines the pollination syndrome of any flower class, they have essentially the same general features over the entire world. Thus convergent evolution for floral structure and habit is a necessary product of any widespread pollination system. The variations in floral habit and structure among the Proteaceae putatively pollinated by nonflying mammals therefore reflect differences in (1) modes of locomotion to the flowers (i.e., terrestrial or arboreal movements as opposed to flying) and (2) foraging behavior. The great mobility of the nonflying mam- mals around flowers and the highly developed chewing apparatus (particularly among generalized feeding rodents) would probably make an attracting unit con- sisting of a single, large flower nonadaptive because of the destructive nature of these animals. The Proteaceae have apparently compensated for the highly destructive activities of these apparent pollinators by increasing the number of reproductive units far beyond what is necessary to maintain successful repro- ductive levels. In the South African cryptic, geoflorous species seed set is con- sistently low, usually below 5%. The same is true in the corresponding Australian genera (A. George, personal communication). In Banksia, the number of flowers per spike probably exceeds 4,000, yet the mature fruits are so large that it would be a physical impossibility for more than perhaps 50 to develop (Fig. 12). In addition to maximizing flower production, the flowering patterns in the heads are staggered temporally so that only several outer whorls are in anthesis simul- taneously. If all the flowers opened concurrently, and in view of their sweetness 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 at anthesis, the entire inflorescence might be more easily destroyed by attracting a large number of pollinators. If generalist feeding, nonflying mammals are potentially so aggressive and opportunistic in exploiting food sources, one might ask why they rarely disturb the typically bird-pollinated South African proteas. First, the bird-pollinated species are not odoriferous and the terminal inflorescences are typically borne high above ground level (Fig. 9); thus most such inflorescences probably escape detection. Another aspect, however, is the presence in some species (especially section Speciosae) of a thick layer of trichomes over the top of the heads. Such a dense layer of trichomes might serve to discourage mammals from chewing to the base of the heads where the nectar is located and thus act as a kind of “mam- mal guard." Birds, of course, easily probe through this layer with their long bills. The heads are also closely surrounded by stiff leathery bracts, which also have considerable trichome development along their margins, where chewing is most apt to be initiated. Excellent illustrations of these phenomena are found in Rous- seau (1970). Finally, the acrid taste of these bracts, as opposed to the sweetness of the bracts and styles in the cryptic, geoflorous species, might also be important. Many observations of nonflying mammals on Proteaceae (and also Myrtaceae) are reported in the popular natural history literature of Australia, e.g., Serventy & Raymond (1974) and Russell (1974) (see also references in Table 2). In fact, so prevalent are these observations that many Australian biologists take this pol- lination system essentially for granted ( Morcombe, 1968; Johnson & Briggs, 1975). To our knowledge, however, no definitive data to establish this relationship has yet been published. Of the nonflying mammals listed in Table 2 as potential pollinators, the honey possum or noolbenger (Tarsipes spencerae), appears to be the best known and apparently the most highly specialized for nectar (and pol- len?) feeding. This amazing animal was studied in captivity by Glauert (1958) and Vose (1972, 1973), who include illustrations. Because no comprehensive re- view of its spectacular adaptations for nectar (and pollen?) feeding is evidently available, a brief resumé of these characteristics taken from the sources quoted above and from Carlquist (1965) might be useful. The head and body are small (6-8 cm long and weighing only 7-11 g). The elongated, tapering snout composes two-thirds of the head. The ears are set far back on the head and the nose is grooved. These features no doubt allow the honey possum to probe deeply into flowers. The tail is longer than both the head and body (8-10 cm) and is prehensile, while the digits of the limbs are slender and elongated for grasping; both characteristics being excellent modifica- tions for the arboreal habit. But it is in the mouth where the most fascinating adaptations for nectar (and pollen?) feeding exist. The tongue is extensible to twice its normal length, tapered, slightly serrated on the margin and brushed at the tip (Fig. 2 in Vose, ). It is exserted through a funnellike structure at the tip of the tapering snout where the lips are modified into flanges. The palate is characterized by ridges which apparently remove accumulated nectar (and pollen?) from the tongue when it is retracted. The jaws are much reduced and dentition rudimentary. Only the upper canines and lower incisors are developed and these appear to function largely in orienting 1977] ROURKE & WIENS—NONFLYING MAMMAL POLLINATION 15 the tongue during retraction. There is no caecum, such a digesting organ for solid food apparently being superfluous in an animal adapted to a nectar diet. Additional structural and especially physiological adaptations for nectar (and pollen?) feeding will no doubt be discovered when more extensive studies are conducted. The evidence that Tarsipes is adapted to a diet derived from flowers (and probably occasional insects) is overwhelming. As a corollary, the conclu- sion that Tarsipes has coevolved as a pollinating agent with various proteaceous (and myrtaceous) genera is inescapable. Sleumer (1955) states that in both northeastern Australia and southeastern New Guinea, the sugar glider (Petaurus breviceps), along with several flower birds, are always associated with flowering Banksia dentata and the myrtaceous genera Melaleuca and Eucalyptus. According to Sleumer, the sugar glider sucks nectar with a “wormshaped” tongue, suggesting possible anatomical adaptations for a nectar diet. Although reference has so far only been made to the cryptic, geoflorous pro- teas, other Australian proteaceous and myrtaceous genera such as Eucalyptus and Melaleuca are known, or suspected, to be visited by various nonflying mammals. For example, Tarsipes reportedly feeds on Hakea and Beaufortia (Morcombe, 1968). Vose (1972) also lists species of Callistemon and Grevillea from which Tarsipes will extract nectar in captivity. Additional reports of nectar sources for flower-visiting marsupials include Dryandra (Glauert, 1958) and Angophora (Porsch, 1934). If the honey possum and possibly also other small arboreal marsupials have apparently coevolved in Australia, why has coevolution between Proteaceae and true rodents not occurred in South Africa? One obvious reason is that the proteas in South Africa ostensibly pollinated by nonflying mammals do not flower throughout the year. The flowering period for these plant groups is limited primarily to late winter or early spring, as previously mentioned. Thus coevolu- tion is impossible because the flowers do not provide a constant food source for these animals which are active throughout the year. Furthermore, it is likely that plants can adapt relatively easily to a generalized feeder, such as many rodents, and that pollination can be reasonably well assured by offering high rewards and reducing competition with other food sources in the community by flowering at the low point in the food cycle. POTENTIAL POLLINATION BY NONFLYING MAMMALS IN OTHER PLANT Groups Discussion in this paper is confined essentially to the possible pollination of Proteaceae by nonflying mammals in South Africa and Australia because we ob- served many of these species and genera in the field. In any overall consideration of the phenomenon, however, other plant groups should not be overlooked. If pollination by small, arboreal marsupials occurs in Australian Proteaceae, it prob- ably also occurs in Myrtaceae. The mouselike lemurs on Madagascar (Sussman & Tattersall, 1976) which take nectar from introduced kapok must be adapted for visiting similar indigenous flowers as well. Porsch (1935) mentions Mada- gascan Symphonia (Guttiferae) as a possible flower adapted for pollination by 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 nonflying mammals. He also discusses other families, e.g., Bombacaceae and Lecythidaceae, which also might have adaptations for pollination by various nonflying mammals. Porsch’s observations merit careful reconsideration, and, especially, critical field studies to test his hypotheses. LITERATURE CITED Bacın, D., B. MULLINS & 8 HunLEy. 1972. A Treasury of Australian Wildflowers. Ure Smith Pty., Ltd., Sydne BENTHAM, G. 1870. c A In Flora Australiensis. Vol. 4-565 BREEDEN, S. & K. BnkEpEN. 1970. Living Marsupials. William 1 (Australia) Ltd., 1972. Australia’s South East. A Natural History of Australia: 2. Taplinger Publ. Co., Inc. New York. CanLQuisr, S. 1965. Island Life. Natural History Press, Garden City, New York. . 1974. Island Biology. Columbia Univ. Press, New Yor Cor, M. J. & F. M. Isaac. 1965. Pollination of the baobab (Adansonia digitata ` ) Be F 8 bush baby ( Galago crassicaudatus E. Geoffroy). E. African Wildlife DEGENEn, O. 1945. Plants of Hawaii National Park Illustrative of = ed pa tes Of y South Seas. Edwards Bros., Inc., Ann Arbor, Michigan. ERICKSON, R., S. GEORGE, N. G. MARCHANT & M. K. MoncoMBE. 1973. Flowers and Plants of 888 Australia. A. H. & A. W. Reed ( Pty.) Ltd., Perth. Farcri, K. & L. VAN DER PIII. 1971. Principles of Pollination Ecology. Ed. 2. Pergamon Press, Ltd., Nu FREELAND, W. J. & H. JANZEN. 1974. Strategies in ud by mammals: the role of plant secondary ni ue Amer. Naturalist 108: 269-289 Guavert, L. 1958. The honey mouse. Austral. Mus. Mag. B 284-2 285. Grant, V. 1950. The protection of the ovules in flowering plants. x ero 4: 179-201. Horn, W. 1962. Breeding research on South African plants: Il. ility of Proteaceae. . S. African Bot. 28: 259-268. aras L. A. S. & B. G. Briccs. 1975. On the Proteaceae—the evolution and classification 1 caris Bot. J. Linn. Soc. 70: 83-182. Lom LEss, A. R. 1961. The nutritional interpretation of sclerophylly based on differences in iii che pons composition of sclerophyllous and mesophytic leaves. Ann. Bot. (London), n.s., 25: —184. ae M. K. 1968. Australia's Western Wildflowers. Landfall Press, Perth. Perren, J. J. 1962. Ecological and behavioral studies of Madagascar lemurs in the field. a. New York Acad. Sci. 102(2): aad: 8 E. P. 1912. Proteaceae. In Flora Capensis. Vol. 5: 560-5 * O. ur Süugetiere als 1 und die Frage a Siiugetierblume. I. iol. Gen. 10: 657-685. Säugetiere als Blumenausbeuter und die Frage der Süugetierblume. II. Biol. 188 . 1936a. Säugetiere als Blumenausbeuter und die Frage der Süugetierblume. HMI. Biol. Gen. 12: 1-21. . 1936b. Süugetierblumen. Forsch. & Fortschr. 12: 207 pig M, & P. Yeo. 1972. The Pollination of Flowers. Taplinge er Publ. Co., New York. Rao, C. 1971. Proteaceae. Botanical Monograph 6. Council of Scientific and aussi mne New Delhi. Raven, P. H. 1972. Why are bird-visited flowers predominently red? Evolution 26: 674. Ripe, W. D. L. 1970. A Guide to the Native Mammals of Australia. Oxford Univ. Press, Melbourne. RonEnTs, A. 1951. In The Mammals of South Africa, R. Bigalke, V. FitzSimons, & E. Malan (editors ). pus by the trustees of the mammals of South Africa book rie) Hafner prr Co., N.Y. RounkE, J. 969. 5 studies on Sorocephalus and Spatalla. J. S. African Bot. Suppl. 7 xs A ROUSSEAU, F. 1970. The Proteaceae of South Africa. Purnell & Sons (S.A.) Pty., Ltd., Cape own. RusseLL, F. 1974. Marsupial mysteries and marvels. Audubon 76: 14-35. 1977] ROURKE & WIENS—NONFLYING MAMMAL POLLINATION 17 SERVENTY, V. & R. RAYMOND. 1974. The honey possum. Austral. Wildlife Heritage. 4(48): 1505-1 509. agen C.J. 1967. The Sunbirds of Southern Africa, Also the Fay, «ome the White-eyes nd the Spotted Creeper. South African Bird Book Fund, A. A. Balkema, Cape Town. sissy H. 1955. Proteaceae. In Flora Malesiana, C. G. G. J. van pe (editor). Ser. 1, 5(2): 147-206. b R. W. & I. TATTERSALL. 1976. Cycles of activity, group composition, and diet of lemur mongoz mongoz Linnaeus 1776 in Madagascar. Folia Primatologia 26: 270—283. Vose, H. 1972. Some ae ai on a honey possum (Tarsipes spencerae) in captivity. W. Austral. Naturalist 12: 61-67. 1973. PT habits of the western Australian honey possum, Tarsipes spencerae. J. Maan): 54: Wirp, H. 1968. E. ds in South Central Africa. Kirkia 6: 197—222. EVOLUTION OF SEED SIZE, SHAPE, AND SURFACE ARCHITECTURE IN THE TRIBE EPILOBIEAE (ONAGRACEAE)! STEVEN R. SkAvEY?, Rogert E. MAGILL? AND PETER H. RAVEN‘ ABSTRACT he seeds of more than half of the approximately 210 species of Onagraceae tribe Epilobieae were examined with the scanning electron microscope. The six species of Boisduvalia s irregularly angular-fusiform seeds with convex, flat, irregularly polygonal surface cells 1 two species and an irregularly striated reticulum formed by the unevenly joining walls of the surface cells in the four others. They are similar to one another and RUE distinct from those of Epilobium, although the relationship between the genera is undoubtedly close. The seeds of Epilobium fall in seven groups: (1) large, obovoid seeds with a more or less prominent micropylar constriction, in three small sections of generalized peel rad ben in one species, E. rigidum, of sect. Epilobium; (2) smaller papillose seeds in over hal the other species; (3) 5 seeds, indepe tidenfiy evolved in many species; (4) 1 patelliform seeds in four Australasian species; (5) irregularly reticulate seeds in one subsection of Epilobium sect. 5 6) ric ged seeds in a ä coherent group of North American origin; (7) finely papillose, distinctive seeds in sect. ssostigma. More or less prominent chalazal beaks have evolved in some species. From xe 1 ic ancestors, Epilobium has evolved a highly successful group of e in sect. Epilobium that have achieved worldwide dis- tribution. This trend seems to have been accompanied by an increase in seed number and a concomitant decrease in very size. The well-marked tribe Epilobieae, one of six that make up the family Onagraceae, includes some 200 species of Epilobium, of worldwide distribution; and six of Boisduvalia, five of western North America, with one common to Argen- tina, and one additional species restricted to western South America. The western North American Zauschneria, often recognized as distinct from Epilobium, is based on a red-flowered, bird-pollinated species of one of the constituent groups of Epilobium. Zauschneria has accordingly been reduced to the status of a section of Epilobium (Raven, 1976). Of the six sections of Epilobium, two, with a total of three species, consist of annuals and are restricted to western North America; two others, with a total of four species, are generalized xerophytic perennials of western North America; one, Chamaenerion, includes seven species of Eurasia, two of which extend to North America; and the remaining one, sect. Epilobium, consists of some 185 species, found on every continent except Antarctica, but especially well represented at high altitudes and high latitudes. The surface sculpturing of the seeds of Epilobium has long been employed as an important taxonomic character (cf. Haussknecht, 1884; Samuelsson, 1923, The authors are grateful to the U.S. National Science Foundation for a series of grants to Peter H. Raven which made these investigations possible. The generous assistance afforded Raven by Mr. W. S. Bertaud, head of the Electron Microscope Section, and Mrs. Lesley A Donaldson. of the Physics and Engineering Laboratory, D.S.I.R., Lower Hutt, New Zeala nd. in carrying out preliminary SEM studies of the seeds of some 75 species of Epilobium in 1969-70 is also gratefully sag dune ? Department of Biology, Lewis & Clark College, eei Oregon 97219. * Botanical Research B stitute, "Private Bag X101, Pretoria, 0001, South Africa * Missouri Botanical Garden, 2345 Tower Grove Avenue, "St Louis, Missouri "63110. ANN. Missoum Bor. Garp. 64: 18-47. 1977 1977] SEAVEY ET AL.—EPILOBIEAE SEEDS 19 1930; Munz, 1965), and it is natural that these seeds have been investigated in recent years with the scanning electron microscope (SEM). The regional or more limited studies that have been concerned with the scanning electron mi- croscopy of the seeds of Epilobium are the following: Berggren (1974), Denford & Karas (1974), Skvortsov & Rusanovitch (1973, 1974), Raven & Raven (1976), and Seavey et al. (1977). To a certain extent, these studies have provided back- ground for the present more comprehensive effort. In addition to the surface variations seen with the SEM, seeds of Epilobicae vary considerably in size and shape. It is the purpose of our report to evaluate these seed characters against a background of other lines of systematic investiga- tion, including cytology, morphology, and biogeography. Included in this analy- sis are more than half of the approximately 210 species of Epilobieae, represent- ing the entire range of diversity; four of the seven species of sect. Chamaenerion; and all species of the remaining sections of Epilobium, as well as all six species of Boisduvalia. The seeds of all but one of the species of Epilobium possess a tuft of trichomes, the coma, on their chalazal (distal) end, whereas none of the species of Boisdu- valia have a coma. Although some variation exists in the color and relative strength of attachment of the coma (cf. Raven & Raven, 1976), no differences in its ultrastructure have been detected and this feature of seed anatomy is not dealt with in the present report. MATERIALS AND METHODS Seeds were collected from recently grown garden plants or from herbarium sheets. The coma was removed in most cases, but left intact if removal damaged the chalazal end of the seed. The seeds were then mounted on aluminum stubs with double-sticky tape, coated with gold/platinum in a Technics Hummer I and examined with a Cambridge Stereoscan Mark 2A scanning electron microscope, operated at an accelerating voltage of 20KV, at the Department of Pathology, Medical School, Washington University. About 75 other samples were photo- graphed in the same way in the Electron Microscope Section of the Physics and Engineering Laboratory, D.S.LR., Lower Hutt, New Zealand, and the results of examining these photographs are likewise incorporated into the present paper. Four to six seeds of each sample were mounted and inspected. Little varia- tion within samples was evident, and the seed photographed was in all cases judged to be typical for the sample. Photographs were taken at approximately 60 x, 240 X, and 1200 x. The higher magnification photographs were taken near the center of the seed, but since little variation is usually evident over the sur- face of this seed, this practice was not strictly followed in all cases. A list of specimens from which seeds were taken for the illustrations (Figs. 1-210) in this paper is presented in Table 1. Voucher numbers are given in the legends of Figs. 1-210 only when more than one collection is listed in Table 1. Bars at the top of each plate indicate, respectively, 0.5 mm, 125 um, and 25 pm. 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE 1. Voucher information for species of Boisduvalia and Epilobium illustrated in this paper. M and G numbers refer to garden planting numbers, used also on herbarium vouchers; R numbers are Raven collections. All vouchers are deposited at the Missouri Botanical Garden ( MO), unless otherwise indicated. Boudica c Tam Curran. U.S.A. Calif. Yolo Co., Crampton 9222 B. Ei i ( Lindl.) S. Wats. U.S.A., Calif., Fresno Co., near Auberry, Seavey in 1974, B. E 5 ( Nutt. ) 8 U.S.A., Calif., Solano Co., near Elmira, Crampton 9219. B. macrantha Heller. „ Calif., Modes Co., near Lookout, Seavey in 1974, M137. B. stricta (A. Gray) 1 U.S.A., Calif., Pues Co., near Auberry, Seavey in 1974, M133. B. subulata (Ruiz & Pavón) Raimann. Chile, Nuble, Chan se d» Watson 4405, 58. Epilobium alpestre (Jacq. ) Krock. Switzerland, Canton Vaud, seed from Conservatoire. et rdin botaniques, Ge neve, 1972-1788, M31. E. Gn Vill. U.S.S.R., y ARN Skvortsov in 1968 (DS), R26179. E. amurense Hausskn. 0 Pref. : saponin Hayakawa-cho, Deguchi in 1974, M80. E. e E PEA Lam. U.S.S.R., E. Chukotka, Iskaten Range, Kozhevnikov in 1972, M7185. E. angustifolium L. subsp. 5 Mosquin, U.S.A., Calif., Onion Valley, Twisselmann 825 (CAS) E. atlanticum Litard. & Maire. Spain, Sierra Nevada, R26166. E. behringianum Hausskn. Alaska, Kiska Quad., Buldir Is., Dick 414, M294. E. billardierianum Sér. subsp. billardierianum. New Zealand, Nelson, Brockie CHR199322, M4 E. canum (Greene) Raven subsp. garrettii (A. Nels.) Raven. U.S.A., Utah, Washington Co., Zion Natl. Park, Seavey in 1974, GS65. E. canum subsp. canum. U.S.A., Calif., Santa Barbara Co., Santa Cruz Is., seed from Rancho Santa Ana Botanic Coden C844. E. canum subsp. septentrionale (Keck) Raven. U.S.A., Calif., Humboldt Co., Trinity River, Tracy 5974. E. canum subsp. latifolium (Keck) Raven. U.S.A., Calif., Lake Co., Snow Mt., Seavey in 2807. E. chilense Hausskn. Chile, Prov. Cautín, near Lake Icalma, Zöllner 7868, M108. E. ciliatum Raf. U.S.A., Oregon, Josephine Co., near O'Brien, Seavey t E. coloratum Biehler. See ds from Copenhagen Botanical Gardens, ASS. E. davuricum Fisch. Canada, Yukon Terr., Porsild 306 (UBC); U.S.S.R., S.E. Chukchi Pe- ninsula, Yurtsev & Raszhivin in 1972 E. denticulatum Ruiz & Pavón. Peru, Pampa to Yamobamba, ca. 70 km E. of Trujillo, Conrad S E. dodonaei Vill. U.S.S.R., no voucher. 7. duriaci Gay ex Godron. n Puerto Ventana, Oviedo, Merxmiiller & Grau 21360, R26258. 2 5 Drew. U.S.A., Calif., Siskiyou Co., Seavey in 1971, M409; Washington, Clallam p. 1111; M559. E. i dH. Lév. Japan, Tottori Pref., Yamamoto in 1970, M 265. E. foliosum (Nutt. ex Torr. & A. Gray) Suksd. U.S.A., Oregon, Douglas Co., Raven 19089. E. glaucum Phil. Chile, Prov. Curicó, Dept. Curicó, Marticorena, Matthei & Rodriguez 1, M68. E. gunnianum Hausskn. Australia, N.S.W., New England Natl. Park, Raven & Englehorn 853 — pA E E. hirsutum L. U.S.S.R., E. Kazakhstan, Altai Mts., Belianina in 1969 (DS), M58. I. m Samuelsson. Peru, 20 km W. of Arequipa, pede tt 1004, M341. i. hornemannii Reichenb. s. lat. U.S.S.R., E. Chukchi Peninsula, Yurtsev & Sutin in 1971. E. komarovianum H. Lév. New Zealand, near Mt. Cook, Raven & Engelhorn. CH R-199430 (MO). E. latifolium L. U.S.S.R., Dist. d Makeeva in 1971. E. leiophyllum Hausskn. Afghanistan, S. of Unai Pass, Breckle A2717, G352. E. luteum Pursh. Alaska, Juneau, Shumway in 1891 (GH). E. minutum Lindl. ex Lehm. U.S.A., Calif., Plumas Co., Howell 51156. 1977] SEAVEY ET AL. 21 TABLE 1. (continued) E. nesophilum Fern. Canada, St. Phillips, Newfoundland et in 1974, E. ines Munz. U.S.A., Nevada, Clark C du cua 1 Mts., 9 in 2 7 6808. z. nivium T. S. Brar ndegee. U.S LA. Ca lif., Lake Ge. Snc wM ts. , Seavey in 1974, G866. ' 5 Seid Czechoslovakia, W. P. Kral in 7967, G E. obcordatum A. Gray subsp. obcordatum. U.S.A., Calif., vm Co., Oneida Lake, Mc- Millan 59-1 (CAS) B. m subsp. siskiyouense Munz. U.S.A., Calif., Siskiyou Co., Sacramento River head- ers, Pringle 1882 (NY). E. P oan Schreb. England, Surrey, Raven 26092, M2. Z. oreganum Greene. U.S.A., Oregon, Josephine Co., near O'Brien, 3 1117. E. oregonense Hausskn. U.S. A., Calif., Mono Co., Ro ak Creek, Seavey in 1970, G350. E. palustre L. U.S.S.R., Chukchi 1 Y itio dr Raszhivin in 1972. E. paniculatum Nutt. ex Torr. & A. Gray. U.S.A. pote Josephine Co., near O'Brien, Seavey n 1975, M554. E. P pauciflorum Samuelsson. Chile, on the pass of 1 5 Zöllner 6245, M110. E. pictum Petrie. New Zealand, Tasman heres Raven & Wilson 25617. E. 5 C. B. Robinson. Taiwan, Hualien Co., K. S. Hsu 1714, G657. E. pylaieanum Fern. Canada, Newfoundland, St. Stephen’s, Olsen in 1974, M127. E. 5 Fr. & Sav. Japan, Nikko, seeds from University of Tokyo Botanical Garden, Tochi, M73. E. rigidum Hausskn. U.S.A., Oregon, Josephine Co., Seavey 1116; Calif., Del Norte Co., Curtis 1 (RSA E. scalare Fern. Canada, Newfoundland, Highlands of St. John, Fernald & Long 28728 (GH). E. shiroumense Matsum. & Nakai. Japan, Pref. Yamanashi, Deguchi in 1974, M81 7. stereophyllum Fresen. Kenya, Aberdare Natl. Park, Raven 26164, M78. 7. stevenii Boiss. Iran, Azerbaijan Prov., Termé in 1971. E. strictum Muhl. Canada, Ottaw Onlar. Jarleton Co., Frankton in 1974, M65. E. suffruticosum Nutt. U.S.A., Wy: oming, Teton Co., Teton Natl. Park, Raven 26464. 7. treleasianum H. Lév. British Columbia, near Boul, Price 1900 (GH). Argentina, Estancia Moat, Tierra del Fuego, Moore 1686, M254; like E. cunninghamii E isskn. except for puliesoeuce OBSERVATIONS I. Epilobium The results are presented according to the sections recognized by Raven (1976). Sect. Cordylophorum (3 species): Epilobium nevadense (Figs. 1-3), E. nivium (Figs. 7-9), and E. suffruticosum ( Figs. 4-6), all have relatively large seeds (2.0-2.7 mm) with a prominent con- striction toward the micropylar end. The seeds of E. nevadense are obovoid, those of E. suffruticosum clavate, and those of E. nivium broadly obovoid. The cells at the point of attachment of the coma form a distinct, although small, neck region at the chalazal end. The surface cells of E. nevadense (Fig. 3) are unique in shape. The center of each cell is occupied by a thick, crater-shaped, apparently collapsed tangential wall. The surface cells of the two other species are entirely convex, giving a cob- blestone appearance at high magnification (Figs. 6, 9). With respect to seed characters, E. suffruticosum resembles E. nevadense more than either resembles E. nivium, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 bo bo FicunEs 1-12. iare electron micrographs of seeds of Epilobium, sects. Cordylopho- rum and Xerolobium. — 3. nevadense 5, —4-6. E. suffruticosum (Cordy- lophorum).—7-9. E. nivium (Cordylophorum).—10-12. E. paniculatum (Xerolobium). 1977] SEAVEY ET AL.—EPILOBIEAE SEEDS 93 Sect. Xerolobium (1 species): The large, broadly obovoid seeds of E. paniculatum (Figs. 10-12) also have a prominent constriction toward the micropylar end. The tangential walls of the surface cells have a centrally situated convex prominence (Fig. 12). The seeds of this species are similar to those of E. nivium. The neck region is incon- spicuous. Sect. Zauschneria (1 species): The seeds of four of the six subspecies of Epilobium canum are presented in Figs. 13-24. Resembling those of the preceding section, these seeds are large and broadly obovoid; and, like all of the species of the preceding two sections, they have a constriction toward the micropylar end. The neck region is inconspicuous, but, as in those of the preceding sections, distinct, especially when viewed from a low angle (e.g., E. canum subsp. canum, Fig. 199). The tangential walls of the surface cells are entirely convex with little radial wall evident, except for subsp. canum (Figs. 16-18), in which the tangential walls are convex but sunken within prominent radial walls. Sect. Chamaenerion (7 species): he seeds of this section are small (1.0-1.8 mm) and relatively narrower than those of the preceding three sections (Figs. 25-36). The neck region is evident in dorsal view and is composed of irregularly constricted chalazal end cells (e.g., E. latifolium, Fig. 209). The surface cells of the three species of subsect. Leiostylae, E. angustifolium (Figs. 25-27), E. conspermum Hausskn., and E. latifolium (Figs. 28-30), lack convex tangential walls, and the major feature of the surface is the irregularly polygonal reticulum formed by the radial walls (Figs. 27, 30). The two species of subsect. Rosmarinifolium we examined, E. stevenii (Figs. 31-33) and E. dodonaei (Figs. 34-36) are of approximately the same size and shape, but the surface tangential walls appear as raised, irregu- larly compressed areas in the center of prominent regularly polygonal radial walls. Sect. Crossostigma (2 species): These small, obovoid seeds, blunt at both ends, are unique in the tribe in their very finely papillose surface cells (Seavey et al., 1 . The surface cells of E. minutum (Figs. 193-195) are concave and very finely papillose over the tan- gential walls as well as the prominent reticulate radial walls (Fig. 195), whereas the surface in E. foliosum (Figs. 196-198) has isolated convex, smooth tangential walls surrounded by finely papillose, elevated radial walls. The neck region in both species is inconspicuous. Sect. Epilobium (ca. 185 species): Extensive variation exists in the size, shape, and surface architecture of the species of this large section. Representatives of the species with the largest seeds, E. rigidum (Figs. 37-39, 40-42), may possess a prominent (Fig. 37) or relatively obscure (Fig. 40) constriction toward the micropylar end, resembling those of sect. Cordylophorum. The surface cells of this species are convex over most of their surface, and the radial walls are mostly obscured (Figs. 39, 42). 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficures 13-24. Scanning electron microgr raphs of seeds of Epilobium sect. Zauschneria. —13-15. E. canum subsp. garrettii.—16— E. canum subsp. canum.—19-21. E. canum subsp. septentrionale.—22-24. E. canum subsp; latifolium. 1977] SEAVEY ET AL.—EPILOBIEAE SEEDS 95 _Ficures 25-36. Scanning electron micrographs of seeds of Epilobium sect. Chamaenerion. 5-27. E. angustifolium.—28-30. E. latifolium.—31-33. E. stevenii.—34-306. E. dodonaei. T first two species belong to subsect. Leiostylae, the second two to subsect. Rosmarinifolium. 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 FIGURES 37-48. Scanning electron micrographs of seeds of Epilobium sect. Epilobium. —37-39. E. rigidum, Seavey 1116.—40—42. E. rigidum, Curtis 1.—43-45. E. obcordatum subsp. obcordatum,—46—48. E. obcordatum subsp. siskiyouense. m= SEAVEY ET AL.—EPILOBIEAE SEEDS 97 1977] Leg 2 „A j E SS Y Ficures 49-60. Scanning electron micrographs of seeds of Epilobium sect. Epilobium. — 49-51. E. shiroumense.—52-54. E. fauriei —55-57. E. platystigmatosum.—58-60. E. stereo- phyllum. 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficures 61-72. Scanning electron micrographs of South American species of Epilobium sect. Epilobium.—61—63. E. cf. pauciflorum.—64-66. E. denticulatum.—67—69. E. glaucum. 70-7 ;. Sp 2. L. sy 1977] SEAVEY ET AL.—EPILOBIEAE SEEDS 29 ae e d dee Ficures 73-84. Scanning electron micrographs of seeds of go on Epilobium. —73-75. E. hirtum.—76-78. E. oregonense.—79-81. E. coloratum.—82-8: . leiophyllum. This combination of characters is unique in sect. Epilobium. The seeds of E. obcordatum (Figs. 43-45, 46-48) are similar in surface relief (esp. Fig. 45), but they are smaller and lack a prominent constriction at the micropylar end. The remaining species of sect. Epilobium have seeds which are smaller, lack 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 a micropylar constriction, and are characterized by surface cells which, although often convex, always have clearly evident radial walls. The range in size of this remaining group of species extends from approxi- mately 0.5 mm long for E. denticulatum (Figs. 64-66), E. komarovianum (Figs. 121-123), and E. pictum (Figs. 124-196) to approximately 1.5 mm for E. cf. pauciflorum (Figs. 61-63), E. pylaieanum (Figs. 88-90), and E. strictum (Figs. 85-87). The range in shape of this group of species extends from the obovoid seeds of E. hirsutum (Figs. 112-114) and E. obscurum (Figs. 109-111) to the narrowly obovoid seeds of E. strictum (Figs. 85-87) and E. shiroumense (Figs. 49-51). In overall shape, E. gunnianum (Figs. 118-120) stands out as having an extended, flat ventral surface which extends beyond the outline of the main body of the seed. In many species the cells in the region of attachment of the coma proliferate to varying degrees, thus forming a distinct neck. This neck is clearly evident in dorsal view in E. alsinifolium (Figs. 145-147), E. anagallidifolium (Figs. 148- 150), E. atlanticum (Figs. 127-129), E. chilense (Figs. 172-174), E. ciliatum (Figs. 169-171), E. denticulatum (Figs. 64-66), E. hirtum (Figs. 73-75), E. hornemannii (Figs. 136-138), E. oreganum (Figs. 160-162), E. oregonense (Figs. 76-78), E. scalare (Figs. 154-156), and E. shiroumense (Figs. 49-51). This neck may exceed 0.2 mm in E. davuricum (Figs. 94-96) and E. pylaieanum (Figs. 88-90). Although the neck may not be seen as a distinct region in some species when viewed from directly above with the seed lying flat, it can be dis- cerned if the seed is tilted (e.g., E. watsonii, Figs. 157, 202). The neck region may be a moderately thick extension of the chalazal end cells (e.g., E. strictum, Fig. 203; E. exaltatum, Fig. 205) or it may be relatively thin, in which case it is displaced toward the ventral side (e.g., E. anagallidifolium, Fig. 204; E. oregonense, Fig. 206; E. scalare, Fig. 207). In species with the longest necks, the neck cells appear individually elongated (e.g., E. davuricum, Fig. 208). The individual trichomes of the coma are usually inserted at the very apex of the neck, although they are occasionally inserted over a broader area (e.g., E. palustre, Fig. 210). If the neck is relatively thin, as in E. exaltatum (Fig. 205) or E. wat- sonii ( Fig. 202), it is sometimes pellucid. There are three types of surface cells in sect. Epilobium: papillose (Group I; Berggren, 1974), cells with a convex portion centrally located on the tangential wall, variously shaped and isolated from its neighboring cells by a prominent radial wall reticulum; foveolate ( Denford & Karas, 1974; Group III; Berggren, 1974), cells lacking any prominent feature other than the raised, regularly poly- gonal radial walls; and ridged (Group II; Berggren, 1974), cells with a centrally located convex portion which is laterally compressed and fused with the raised portions of neighboring cells, forming longitudinal ridges along the length of the seed. Most species of sect. Epilobium have papillose seeds. The central papilla of the surface cells of many species is irregularly compressed into a multisided prominence as in E. alpestre (Figs. 103-105), E. amurense (Figs. 97-99), E. obscurum (Figs. 109-111), E. shiroumense (Figs. 49-51), and E. stereophyllum 1977] SEAVEY ET AL.—EPILOBIEAE SEEDS 3] q n „ %% 9 rege e. (9 EA FicunEs 85-96. Scanning electron „ E i of Epilobium sect. Epilobium. — 85-87. E. strictum. — 8-90. E. pita pen .—91- E. davuricum, Porsild 306.—94—96. 972. E E. dav uricum, Yurtsev d» Rasz hivin in 19 39 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 100 ; FiGungs 97-108. Scanning electron micrographs of seeds of Eurasian species of Epilobium sect. Epilobium.—97-99. E. amurense.—100-102. E. pyrricholophum.—103-105. E. alpestre. —]106-108. E. duriaei. 1977 SEAVEY ET AL.—EPILOBIEAE SEEDS 33 Ficures 109-120. Scanning electron micrographs of se 15 of Epilobium sect. Epilobium. — 109-111. E. obscurum.—112-114. E. hirsutum.—115-117. E. billardierianum subsp. billardierianum.—118-120. E. gunnianum. 34 FIG URES 121-13 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 . Scanning electron micrographs of E b Epilobium sect. 5 121-12 E. ene ianum.—124-126. E. pictum.—127-129, E. atlanticum.—130—13 "nein 1977] SEAVEY ET AL.—EPILOBIEAE SEEDS 35 FIGURES Se 144. Scanning electron ane e of seeds of E erue sect. Epilobium. —]133-135. E. behringianum.—136-138. E. hornemannii s. lat. —139— . luteum.—142- 144. E. tre * 'asianum. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficures 145-156. Scanning electron micrographs of seeds of Epilobium sect. Epilobium. 7 ; 3. E —145-147. E. alsinifolium.—148-150. E. 154-156. E. scalare. anagallidifolium.—151—153. E. nesophilum.— 1977] SEAVEY ET AL.—EPILOBIEAE SEEDS 37 n RE yy 1 ay ia - ERA oe Ficures 157-168. Scanning electron micrographs of se GA X North American Vae of Epilobium sect. Epilobium.—157-159. E. watsonii.—160— E. oreganum.—163— E. exaltatum, M409.—166—168. E. exaltatum, M559. 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 FIGURES . 74. eee: electron i raphs of seeds of Epilobium sect. Epilobium. 39-171. E. 1 172-174. E. chilen (Figs. 58-60), whereas in others it remains more Toe domeshaped, as in H. denticulatum (Figs. 64-66), E. fauriei ( Figs. 52-54), E. glaucum (Figs. 67-69), and E. pyrricholophum ( Figs. 100-102). In some species, the papilla is marked by radial lines which appear as fine ridges, as in E. oregonense (Figs. 76-78), E. Agen anum (Figs. 88-90), some samples of E. billardierianum (Figs. 115-117), 2. davuricum (Figs. 91-93), E. hirsutum (Figs. 112-114), and E. nesophilum s 151-153). The papilla of some species, such as E. leiophyllum (Figs. 82-84) and E. duriaei (Figs. 106-108), is collapsed in the center. The surface cells are usually arranged in irregular rows running longitudinally the length of the seed, but in some species, such as E. amurense (Figs. 97-99), E. fauriei (Figs. 52-54), and E. obscurum (Figs. 109-111), the rows are distinctly regularly ar- ranged. In foveolate seeds the regularly polygonal reticulum formed by the radial walls is the most prominent feature of the seed surface. This radial wall reticulum is relatively low on the seeds of E. gunnianum ( Figs. 118-120), and E. komarovia- num (Figs. 121-123), but more pronounced on the seeds of E. alsinifolium (Figs. 146-147), E. anagallidifolium (Figs. 148-150), E. atlanticum (Figs. 127-129), E. behringianum (Figs. 133-135), E. hornemannii (Figs. 136-138), E. luteum ( Figs. 139-141), E. nutans (Figs. 130-132), and E. pictum (Figs. 124-126). Ridged seeds, marked by longitudinal rows of laterally compressed, fused ridges, are found principally in North American species. Among these are E. ciliatum (Figs. 169-171), E. exaltatum (Figs. 163-165), E. oreganum (Figs. 160-162), and E. watsonii (Figs. 157-159). The South American E. chilense — 1977] SEAVEY ET AL.—EPILOBIEAE SEEDS 39 “1GURES 175-186. Scanning electron micrographs of seeds of the four species of Bois- duvalia sect. Boisduvalia.—175-177. B. densiflora.—178-180. B. macrantha.—181—183. B. stricta. —184-186. B. subulata. 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 — FIGURES 187— Scanning e d micrographs of seeds of the species of Bc 7 aq sect. Currania 28 j^ nilobium sect. Crossostigma.—187-189. Boisduv d cleistogama.—19% 192. B. glabella— 193-195. E pilabium n 1 196-198. E. foliosu- 1977 SEAVEY ET AL.—EPILOBIEAE SEEDS 4] Figures 199-210. Scanning electron micrographs showing variation in 5 E. A] of seeds of Epilobium, including the dev Roue of more or less pronounced bea tax are members of sect. Epilobium except as indicated in 1 ama following cheir names. — 199. Epilobium canum d canum (Zauschneria) —200. E. rigidum, Curtis 1.—201. E. obcordatum subsp. obcordatum.—202. vat: 0 2 Pc. uer M4. E. anagallidi- folium.—205. E. exaltatum. — 906. E. 97 nse.—207. E. scalare.—208. E. davuricum, Yurtsev & Raszhivin in 1972.—909. E. 1 m —210. 55 palustre. 42 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 (Figs. 172-174) likewise has ridged seeds. The seeds of hybrids between species with ridged and papillose seeds are intermediate in various ways and have been illustrated by Skvortsov & Rusanovich (1973). Il. Boisduvalia The seeds of Boisduvalia are distinct from those of Epilobium. Their shape, which is irregularly angular-fusiform, is unique, as is the structure of their sur- face cells. The relatively broad seeds of B. densiflora (Figs. 175-177) and B. subulata (Figs. 184-196) have surface cells which are slightly raised, flat, and irregularly polygonal. The seeds of B. macrantha (Figs. 178-180), B. stricta (Figs. 181-183), and the narrower seeds of B. cleistogama (Figs. 187- 189) and B. glabella (Figs. 190-192), have surfaces that are covered by an ir- regularly striated reticulum composed of the unevenly joining radial walls of the surface cells. DISCUSSION A number of distinct seed types occur in the genera Boisduvalia and Epilo- bium, as will be discussed below. In both genera, as in other Onagraceae, the surface architecture is made up of the regularly repeated structures of individual surface cells. The characteristic forms of these surface cells are the result of dif- ferential thickening in their walls, as thin sections of the seeds of Epilobium (Ky- tovuori, 1972; Denford & Karas, 1974) demonstrate; seed coat patterns in many other genera are comparable, for example in Cordylanthus (Chuang & Heckard, 1972), Melastomataceae (Whiffin & Tomb, 1972), and Mentzelia (Hill, 1976), as well as other investigations reviewed by Brisson & Peterson (1976). There is a major contrast in Epilobieae between the angular-prismatic seeds of Boisduvalia, which lack a coma, and the regularly obovoid, flattened ones of Epilobium, which have a coma in all taxa except in a few in which it has been lost. In addition, the surface cells in Boisduvalia are irregular and thus unlike any found in Epilobium. Those of B. densiflora (n= 10) and B. subulata (n= 19) are low, flat, and irregularly polygonal, whereas those of B. macrantha (n= 10) and its probable aneuploid derivative B. stricta (n=9; Raven & Moore, 1965) are concave and have radial walls that are longitudinally striate and ir- regularly thickened. These four species comprise sect. Boisduvalia (Raven, 1976); the remaining two species, which comprise sect. Currania (n= 15), have seeds that resemble those of B. macrantha and B. stricta, although they are smaller. In Epilobium, it is convenient to recognize seven distinct types of seeds, and these will be discussed in turn. 1. Large, obovoid, constricted. In sects. Cordylophorum, Xerolobium, and Zauschneria, the seeds are obovoid, large, and more or less prominently con- stricted toward the micropylar end. Their surface is made up of cells which are smooth and convex, with obscure lateral walls (Figs. 1-24). The relatively prominent side walls in our preparation of E. canum subsp. canum probably are associated with shrivelling. The five species that comprise these three sections are 1977] SEAVEY ET AL.-EPILOBIEAE SEEDS 43 relictual (Raven, 1976), and this seed type is almost certainly primitive for Epilo- bium. In sect. Epilobium, with some 185 species and a worldwide distribution, it occurs only in Ë. rigidum (Figs. 37-39, 40-42), a large-flowered, xerophytic species of the Siskiyou Mountains of northwestern California and adjacent Ore- gon. The Siskiyou Mountains are a well known relict area (Whittaker, 1960), and E. rigidum very likely resembles the basic stock from which the remaining species of sect. Epilobium have diverged. As in E. nevadense, E. nivium, and E. paniculatum, the subtending bracts in E. rigidum are fused to their pedicel in all but the lowermost flowers. The phylogenetic significance of this observation remains to be determined. 2. Papillose. Most species of sect. Epilobium have seeds that are smaller, papillose, obovoid to narrowly obovoid, and lack a micropylar constriction. The lateral walls of their surface cells are prominent. These include E. obcordatum ( Figs. 43-45, 46-48), the species most closelv related to E. rigidum and, like it, a large-flowered xerophyte of the western United States. In addition, the four species of sect. Chamaenerion subsect. Rosmarinifolium, exemplified by E. dodonaei (Figs. 34-36) and E. stevenii (Figs. 31-33), have this seed type. It seems clearly to have evolved from the first type and to have given rise in turn to all of the other more specialized seed types within the genus, with the probable exception of that found in sect. Crossostigma. The tribe Epilobicae seems to have consisted initially of xerophytes, from which the more widespread and numerous mesophytic and hydrophytic species were derived. Perhaps an evo- lutionary trend toward more numerous, smaller seeds accompanied the exploita- tion of such habitats. Seeds of this type characterize more than a hundred species, including the European E. alpestre, E. collinum, E. duriaei, E. hirsutum, E. lanceolatum, E. montanum, E. nervosum, E. obscurum, E. parviflorum, E. roseum, and E. tetrago- num, as well as the circumboreal E. davuricum and E. palustre (Skvortsov & Rusanovitch, 1974: Berggren, 1974; this paper). Papillose seeds also occur in the Asian E. amurense, (Figs. 97-99), E. fauriei (Figs. 52-54), E. platystigmatosum (Figs. 55-57) and E. pyrricholophum (Figs. 100-102): the South American E. denticulatum (Figs. 64-66), E. glaucum (Figs. 67-69), E. hirtum (Figs. 73-75), E. cf. pauciflorum (Figs. 61-63), and one unidentified species (South American species # 1, Figs. 70-72); the African E. stereophyllum ( Figs. 58-60); the Austra- lasian E. billardierianum (Figs. 115-117); and the North American E. coloratum (Figs. 79-81), E. oregonense (Figs. 76-78), E. pylaieanum (Figs. 88-90), and E. strictum (Figs. 55-87), as well as most populations of E. saximontanum Hausskn. Although only one Australasian species of this first type of surface structure is illustrated in this paper, most species of this region have seeds of this type ( Raven & Raven, 1976). Many of the microstructural details of papillose seeds are strikingly constant from one sample to another. The convex portion of the surface cells of E. hirsu- tum (Figs. 112-114), for example, are characterized by spirally arranged radial lines, as illustrated by Skvortsov & Rusanovitch (1974; fig. 1C). Similarly, the surface cells of E. nesophilum (Figs. 151-153), and the closely related E. pylaie- anum (Figs. 88-90) and E. davuricum (Figs. 91-93) have characteristically 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 low convex tangential walls with radial lines as illustrated in Skvortsov & Rusano- vitch (1974: figs. 2E, F, and fig. 2G, respectively), although considerable vari- ability in surface structure within these two species is also evident in both studies (e.g., E. davuricum, Figs. 91-93, 94-96) Nearly all of the species with papillose seeds in sect. Epilobium, well over a hundred, have the BB chromosome arrangement (Seavey & Raven, 1976; un- published). Among the exceptions are the Asian E. fauriei, E. platystigmatosum, and E. shiroumense, as well as most populations of the circumboreal E. horneman- nii s. lat. and the North American E. clavatum Trel., all of which have the CC arrangement. In addition, some of the species with the AA chromosome arrange- ment, including the South American species of the group Denticulata (Samuels- son, 1923, 1930) and the North American E. glaberrimum Barbey and E. brevisty- lum Barbey, as well as E. saximontanum, also have papillose seeds. Both the AA and CC chromosome arrangements differ from BB by one reciprocal transloca- tion, and we believe that each may be independently derived from BB. On the basis of the evidence presented here, it appears that the common ancestor of cach of these groups had papillose seeds. Most species of Haussknecht’s (1884) group Palustriformes, including E. davuricum (Figs. 91-93, 94-96, 208), E. palustre (Fig. 210), and E. pylaieanum (Figs. 88-90) among those illustrated in the present study, have an exaggerated beak at the chalazal end of the seed; although others, such as E. strictum, which undoubtedly belongs to this group, do not (Figs. 85-87, 203). The Palustri- formes have the BB chromosome arrangement and papillose seeds in all taxa; similar in these respects is E. oregonense (Figs. 76-78). Epilobium scalare, a Newfoundland endemic that has been collected only once, has papillose seeds and a prominent beak also but does not resemble Palustriformes in most respects. Chalazal beaks are discussed in general on p. 30 and illustrated in Figs. 199-210. 3. Foveolate. By flattening of the central papilla found in the surface cells of the seeds of the preceding group, foveolate seeds (Danford & Karas, 1974) have evolved. Such seeds, as viewed with a 20x lens, have usually been de- scribed as “smooth” in taxonomic papers on Epilobium. Some species, including the New Zealand E. alsinoides (different subspecies), have some populations with papillose seeds and others with foveolate seeds. The same occurs in the circumboreal E. hornemannii s. lat. and in the North American E. clavatum. It appears, therefore, that the evolution of foveolate seeds has taken place repeatedly within the genus. In Australasia, where all species have the BB chromosome arrangement and presumably evolved from a common ancestor ( Raven & Raven, 1976), 31 species have papillose seeds; 10, including E. komarovianum (Figs. 121-123) and E. pic- tum (Figs. 124-196) have foveolate seeds; E. alsinoides, already mentioned, has both papillose and foveolate seeds in different subspecies; and 4 species, includ- ing E. gunnianum (Figs. 118-190) have a distinctive seed type that will be dis- cussed below. In almost every one of the 11 species in which foveolate seeds oc- cur, these appear to have evolved separately from papillose seeds (Raven & Raven, 1976). In North America, foveolate seeds occur in Epilobium luteum, a rather iso- 1977] SEAVEY ET AL.—EPILOBIEAE SEEDS 45 lated species, and in some populations of the E. glandulosum complex (AA). In addition, they are shanki q qa of some or all populations of several entities in the CC group: E. anagallidifolium (Figs. 148-150), E. behringianum (Figs. 133-135), E. clavatum, and E. hornemannii s. lat. (Figs. 136-138). The seeds of E. treleasianum may be papillose (Figs. 142-144) or foveolate, owing to the hybrid nature of E. treleasianum, a series of populations of hybrid origin between ?. luteum (seeds foveolate) and other species (Seavey, in preparation). Papil- lose seeds appear to be dominant in a genetic sense over foveolate ones. I£urope, only species of the CC ( Alpinac ) Among the species that occur in E. atlanti- group and three other species—E. nutans (BB in part; Figs. 130-132), cum (AA; Figs. 127-129), and E. alsinifolium (AA; Figs. 145-147 )—have fove- olate seeds. Judged from the wide morphological gap between the latter two and the different chromosome arrangement in the first, it is likely that The fact that all three are species group Alpinae E. alsini- species all evolved foveolate seeds independently. of low stature that occur in alpine habitats, like the species of (here defined to comprise only CC species and therefore to exclude folium), suggests that some common selective force may favor foveolate seeds under such conditions. The third European species with the AA chromosome E. alpestre ( Figs. 103-105), has papillose seeds. arrangement, 4. Obovoid-patelliform. Four Australasian species—E. gunnianum (Figs. 115-120), E. curtisiae Raven, E. willisii Raven & Engelhorn, and E. angustum (Cheesem.) Raven & Engelhorn—have distinctive seeds with a hollow ring around their adaxial side, which are thus patelliform. The first three are prob- ably closely related, but the occurrence of similar seeds E. angustum, ap- parently closely related to E. komarovianum, has not been explained satisfactorily ( Raven & Raven, 1976). The seeds of E. willisii and of many populations of E. gunnianum are finely papillose, those of the other taxa foveolate. The obovoid- patelliform seed type undoubtedly evolved from the more frequent papillose type within Australasia. 5. Irregularly reticulate. Leiostylae have seeds in which the very thin radial walls of the epidermal cells of the seed coat form an irregularly polygonal reticulum (Figs. 25-30). The four other species of sect. Chamaenerion, comprising subsect. Rosmarinifolium have papillose seeds that resemble those of most species of sect. Epilobium in size and shape. The close relationship of the two subsections of sect. Chamaene- rion is beyond question in that they share the following unique or highly unusual all leaves spirally arranged, flowers zygomorphic. This indicates unequivocally that the (1974) The three species of sect. Chamaenerion subsect. characteristics for the genus: floral tube obsolete, pollen shed singly. unusual seeds of subsect. Leiostylae, stressed by Skvortsov & Rusanovitch ( and by Brisson & Peterson (1976) as an argument for the generic distinctness of sect. Chamaenerion, represent instead an evolutionary specialization within this group, otherwise characterized by seeds similar to those of many species of sect Epilobium. Also implied by the seed morphology of subsect. Rosmarinifolium is the divergence of sect. Chamaenerion from an ancestor that would be placed within sect. Epilobium long after the divergence of sects. Cordylophorum, Xerolo- bium, and Zauschneria from species such as E. rigidum. Additional evidence for 46 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 the close relationship of sect. Chamaenerion with sect. Epilobium is summarized by Raven (1976) 6. Ridged. Within the North American group of species with the AA chromosome arrangement (Seavey & Raven, 1976), there has originated a dis- tinctive seed type, described above, that doubtless delimits a phylogenetically coherent group of taxa. Illustrated here are E. ciliatum (Figs. 169-171; extends to Japan), E. exaltatum (Figs. 163-165), E. oreganum (Figs. 160-162), and E. watsonii (Figs. 157-159). Epilobium ciliatum occurs as an adventive in Europe and in Australasia ( Raven & Raven, 1976), and its seeds have been illustrated several times (as E. adenocaulon Hausskn., Troughton & Donaldson, 1972: pls. 103-104; Skvortsov & Rusanovitch, 1973, 1974; Berggren, 1974; Raven & Raven, 1976). Virtually identical seeds occur in the South American E. chilense (Figs. 172-174), this suggesting recent immigration to South America following the origin of the group in North America. Within the Epilobium glandulosum com- plex foveolate seeds also occur, but these have probably been derived from ridged ones. The most primitive species of this group is apparently a local en- demic of bogs in the Siskiyou Mountains, E. oreganum, which has exserted, deeply 4-lobed stigmas. Denford & Karas (1974) have interpreted the ridges in seeds of this type as being formed of rows of flattened papillose cells flanked on both sides by foveo- late cells. Our SEM observations, however, show clearly that the ridges are in- stead formed of the finlike central portions of individual surface cells, all such cells on the abaxial side of the seed being similar. No foveolate cells were ob- served on the abaxial surface of these seeds. 7. Finely papillose. The distinctive seeds of sect. Crossostigma, described on p. 23 and by Seavey et al. (1977), cannot easily be related to any other seed type in the genus, and the relationships of the two annual species included in this section are obscure. LITERATURE CITED us ed G. 74. Seed morphology of some Epilobium species in Scandanavia. Svensk Bot. cu es 164—168. Brisson, J. D L. Pererson. 1976. A critical review of the use of scanning electron 5 in “the study of the seed coat. Proc. Workshop on Pl. Sci. Applications of the SEM IIT Res. Inst. 1976 (Pt. VII): 477—495 T. & L. R. HEckanp. 1972. Seed coat morphology in Cordylanthus ( Scrophulari- ceae ) and its taxonomic significance. Amer. J. Bot. 59: 258-265. oues K. E. & I. Karas. 1974. Some os on the seed coat structure within the genus E pilobium. E datus 30: 1144-1 188: HAUSSKNECHT, C. . Monographie der Cutting Epilobium. Jena, Germany. viii + 318 pp. + 23 pls. Hitt, R. J. 1976. Taxonomic and deol on significance of seed coat microsculpturing in Mentzelia ee in Wyoming and adjacent western states. Brittonia 6-112. Kvróvvonr, L 1972. The Alpinae 1 the genus 5 in 5 Fenno— scandia. A morphological, 1 nical, ie 1 e study. Bot. Fenn. 9: 163-203. Mns P.A. 1965. Onagraceae. North Amer. Fl. I : 1-278. Raven, P. H. 1976 (1977). Generic 23 uu Sidon in Onagraceae, tribe Epilo- bieae. Ann. Missouri Bot. Gard. 63: — & D. M. Moore. 1965. A revision of Boisduvalia (Onagraceae). Brittonia 17: 2238-254. & T. E. Raven. 1976. The genus Epilobium (Onagraceae) in Australasia: A 1977 SEAVEY ET AL.—EPILOBIEAE SEEDS 47 systematic and evolutionary study. New Zealand Dept. Sci. Industr. Res. Bull. 216: 1- SAMUELSSON, G. 1923. ripe Sy der sudamerikanischen | Epilobium-Arten. Svensk Bot. Tidskr. 17: 241-296, tab. II-V ra Zur E piobiun: Flora Südamerikas. Svensk Bot. Tidskr. 1-11. Seavey, S. R. R C il evolution in k e sect. Epilobium iis Pl. Syst. Evol, 24: 6- „P. Wi H. RAVEN. t A comparison of Epilobium minutum and E. foliosum. Madrono Cin press ). SKVORTSOV, & I. I. Rusanovircu. 1973. Hybrid cell structures of the seed coat of E pilobium. Priroda (Moscow) 12: 89-93. [In Russian. ] & ———._ 1974. 7 electron microscopy of the seed-coat surface in Epilobium species. Bot, Not. 127: 392-4 TROUGHTON, A. e e 1972. Probing Plant Structure. Chapman & Hall, London. m p WHIFFIN, T. & A. 8. Toms. 1972. The systematic significance y ie cer in the 1 capsular fruited Melastomataceae. Amer. J. Bot. 59: WHITTAKE 960. Vegetation of the Siskiyou wo 1 and California. Ecol. Mosa 30: 279-338. THE BIOSYSTEMATICS OF CALYLOPHUS (ONAGRACEAE )' Howard F. TowwvEn* ABSTRACT The genus Calylophus (Onagraceae), a segregate of Oenothera, was gros in ene ystematic relationships, breeding systems, geno and cytology. re 5 four in sect. Salpingia: C. tubicula, C. hartwegii, C. toumeyi and C. lovanduhfolius. and two in sect. Calylophus: C. berlandieri (formerly C. 3 and C. serrulatus. Several changes in 5 and rank are made. Crosses performed among species demonstrated strong barriers to hybridization 1 n the two sections of the genus and * à moderate barriers among species within sectic pulations of Calylophus are distributed ciunt the Great Plains, the southwestern ale d "tates. and northern Mexico. The various taxa occupy distinct habitats which range from xeric sites in the Chihuahuan Desert to relatively mesic pine and pine-oak forests. most forms of the genus, the plants are suffrutescent perennials and occupy calcareous soils. One form is restricted primarily to gypsum soils lytological investigations showed a remarkable degree of translocation heterozygosity in natural populations of Calylophus. Translocations were observed in all taxa, with 75% of 183 plants (excluding C. Macr nie exhibiting heterozygosity or at least one idi ue Numerous plants were relem ygous for more than one translocation, and the mean num per plant was 1.3. Calylophus serrulatus is a ale seep heterozygote and all indiv a observed were heterozygous for at least five or six translocations. Hybridization Ca wakaqa with € eae ri 5 that C. serrulatus 1 N 5 i hybridity with gametophytic lethals in pollen and embryo sacs vasic AE number of the genus is x = 7. Tetraploidy was observed in individuals from 5 of 62 populations of C. hartwegii m were examined. A few plants of several taxa possessed diminutive chromosomes ranging from 1 to 11 in number. The most frequent observation was of a single dark-staining pair in addition to the normal complement. Chromosome observations of hybrids showed uie found intersectional differences in ructure, primarily from translocations. Translocation differences are a so marked among population in sect. Calylophus, but are slight among de taxa of sect. Salping The breeding systems of Calylophus are varied, with C. serrulatus self. compatible and highly M and the other species self-sterile. bus e berlandieri and C. tubicula have sho oral tubes, strong ultraviolet contrast patterns, matinal anthesis, and are ud yy av mid of cna: insects. dr esis of the remaining members of sect. Salpingia occurs in the afternoon or evening. These plants possess long floral tubes, variable ultraviolet contrast patterns, and are visited by sphingid moths and 1 bees in numbers that vary from locality to locality. Biosystematic studies of the genus Calylophus were begun in 1967, shortly after it had become apparent that this genus constituted a natural group no less distinct from Oenothera than from other genera of the tribe Onagreae (Raven, 1964). This paper is based on those studies and on an examination of extensive ! This study was initiated as a doctoral dissertation at Stanford University ( Towner, 1970b), and subsequently amplified. I would like to express great appreciation to Peter H. Raven, at whose suggestion this research was begun. His advice and generous assistance have been invaluable throughout the course of the study. He, D. E. Breedlove, and D. P. Gregory provided both material of Calylophus and unpublished information on pollination. | Sharon Stewart gave me indispensable assistance in the field, laboratory, and greenhous Dan Holmes and Judith L m cis me in the field. Steven and Ann Seavey were ae helpful in the maintenance of cultivated plants and the handling of herbarium material. John H. Thomas directed the processing of ae loans. The illustrations of flowers were drawn by Julie Spranza The following individuals kindly dae Dr. Raven and me with n 2i Calylophus or with information regarding localities C. Anderson, D. S. Correll, W. 5mery, Ann. Missouni Bor. Garp. 64: 48-120. 1977. 1977 TOW NER—CALYLOPHUS 49 herbarium material. A preliminary publication stemming from this research (Towner & Raven, 1970) was based on a less complete study of herbarium ma- terial and some of the taxonomic decisions made then are changed in the present paper. The genus Calylophus is distinguished from the other genera of the Onagreae by the following suite of characteristics: a peltate stigma which may be discoid or nearly square in shape, sometimes with 4 shallow, broad lobes; microsporog- enous tissue divided into packets in the locules of the anthers; yellow flowers; and a many-seeded capsule. In the opinion of Raven (1964), Calylophus is most closely related to those genera of the Onagreae which share the feature of di- vided microsporogenous tissue: ie, Gaura, Clarkia, Heterogaura, and perhaps Hauya. The genus occurs over much of the Great Plains, extending into the mountains of the Great Basin region and other parts of the Southwest, and also reaching southward to the Mexican Plateau. The area of greatest diversity for the genus includes West Texas, southern New Mexico, and north-central Mexico. Populations are usually colonial and widely scattered, often occurring in dis- turbed areas. Habitats occupied by the species of Calylophus are typically some- what xeric plains or hills with soil that is often calcareous. Plant associations in which the various forms occur range from creosote bush scrub in the Chihuahuan Desert to pine forests of several types. The history of the genus Calylophus has involved several transfers at the generic level as various authors have seen fit to separate the group from Oenothera or to unite the two genera. Traditional treatments of the species here considered have placed them in Oenothera subgenera Salpingia and Calylophus or in the respective genera Galpinsia and Meriolix. Rafinesque (1819) was the first to distinguish a species of what is now Calylophus from Oenothera, although his publication, lacking a description of the genus, was invalid. The name Meriolix was validated only later, by Endlicher, in 1840. Calylophus, the first generic name of legitimate publication, was presented by Spach (1835a) and emended by him to “Calylophis” without justification in the same year (Spach, 1835b). It thus has priority over Meriolix, Galpinsia, and Salpingia at the generic level. Most of the nineteenth century treatments of the species of Calylophus retained them in Oenothera. In the late nineteenth century and until quite recently, the generic names Meriolix and Galpinsia were frequently used for the various species of Ç Pata in treatments such as those of Heller, Small, and Rydberg. — Fisher, R. C. Jackson, W. M. Klein, R. L. McGregor, H. Marshall, T. Mosquin, D. R. ae and the late L. G. Shinners. Pollinating insects m e been most macie l. identified by W. „ J. Burns, H. V. Daly, G. Eickwort, anc Thorp. I am grateful to the curators of the fa hak harkari, from which ES tie tae was secured for the study, or which were visited by the author: ARIZ, CAN, CAS, COLO, DAO, DS, F, GH, K, KANU, KSC, LA, LL, MO, NEB, NMC, NY, OKL, OKLA, PH, 77 "M RSA, SMU, TEN, UC, UNM, US, WTU Financial support during the course of this work was supplied by National Institutes of Health pre puso and heiss FO: fellowships awarded to me and by research grants to eter H. Raven from vM National Science Foundation (1966-1970). Laboratory space and 5 facilities were provided by the Department of Biological Sciences, Stanford Univ I of Biology, Loyola Marymount University, Loyola Boulevard at West 80th Street, Los Angeles, California 90045. 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 During the nineteenth century the explorations and collecting of Nuttall, Wright, Lindheimer, Berlandier, Hartweg, Fendler, James, Cockerell, and Toumey provided type material for most of the presently known taxa in the genus. All taxa now considered valid, except Calylophus hartwegii subsp. mac- cartii and C. tubicula subsp. strigulosus, were described and published between 1817 and 1898. Over this period and during the early twentieth century a num- ber of names were proposed for minor variants, especially by Léveillé, Small, and Nelson. These contributed additional confusion to the taxonomy of the genus, which in any circumstance would have been difficult to interpret. The situation was greatly improved with the publication of Munz's (1929) revision of these species, a treatment which brought together the information available at that time and gave relative order to the taxonomy of the group. Many superfluous names were reduced to synonymy and for the first time decisions were based on adequate herbarium material. Some of the species boundaries remained unclear even then because of the complexity of variation in sect. Salpingia, an erroneous appraisal of the type of C. hartwegii, and the absence of information on breeding systems in the C. serrulatus group. Raven's (1964) paper, which brought this group of species together as the genus Calylophus, was closely followed by the publication of Shinners (1964). Shinners presented a taxonomy similar to that used here, except that the facts concerning breeding systems in C. serrulatus were not yet known, and that some differences in the ranking of taxa exist between our treatments. Other differ- ences include my recognition of C. hartwegii subsp. fendleri and a revised inter- pretation of C. hartwegii subspp. hartwegii and filifolius. In 1965 Munz pub- lished a monograph of the North American Onagraceae (Munz, 1965) in which the species of Calylophus were referred back to Oenothera and in which specific status was granted without justification to several of the entities presented by Munz as subspecies in 1929. Otherwise the taxonomy remained the same as in the earlier publication. A brief taxonomic treatment of this group by Towner & Raven (1970), the basis of my account for the Manual of the Vascular Plants of Texas (Towner, 1970a), considered the species as belonging to Calylophus. The present work generally retains the same boundaries between taxa used in that paper. Excep- tions include two changes in rank, the recognition of C. tubicula subsp. strigulo- sus, the reduction of C. australis to synonymy with C. serrulatus, and the changes in the names for the outcrossing members of sect. Calylophus. All of the fore- going changes were found necessary after a more thorough study of herbarium material was completed. Calylophus lavandulifolius and C. toumeyi were de- termined to be well differentiated from C. hartwegii and thus deserving of specific rank. Calylophus serrulatus is here judged to be possibly of multiple origins, and the series of populations once referred to as C. australis is now thought to be distinguished from other such series only by its geographical separation from the range of the rest of the species. Lastly, C. berlandieri was found to be the ap- propriate name for the entity treated by Towner and Raven as C. drummondianus. Type material of Drummond's was determined by an examination of pollen fer- tility to belong with the complex structural heterozygote C. serrulatus. 1977 TOWNER—CALYLOPHUS 51 CYTOLOGY Mciotic chromosome configurations were determined to establish the frequen- cies of structural and numerical changes in natural populations of Calylophus. Further, the cytology of experimental hybrids was observed in order to describe chromosomal differentiation among taxa in the genus. Previous reports of cy- tology in Calylophus (as species of Oenothera) include those of Hagen (1950), Lewis et al. (1958), Gregory & Klein (1960), and Kurabayashi et al. (1962). These authors reported translocation heterozygosity, tetraploidy, and extra chromosomes. P. H. Raven (personal communication) later established the pres- ence of complex structural heterozygosity in the genus. Those phenomena were confirmed and their taxonomic distribution described in the present study. In- version differences among populations were also found. Table 1 and the sys- tematic accounts of the various taxa combine my findings and the results of earlier investigations. Translocation heterozygosity is extremely common in the genus and was found in all taxa. Calylophus serrulatus, with its system of complex structural heterozygosity, consistently forms rings of 12 or 14 chromosomes at meiotic metaphase I. All other species form mutivalent associations regularly. Excepting tetraploids and C. serrulatus, 130 of 183 plants (71%) had visible translocations, and in this sample the mean number per plant was 1.3. The most frequent num- ber of translocations per plant was 1, but individuals with 2 or 3 were com- mon. In one observation from C. hartwegii subsp. filifolius and two from C. berlandieri subsp. berlandieri, plants were heterozygous for 5 translocations. Directed alternate disjunction seems to prevail since translocation heterozygosity does not appreciably lower pollen fertility in natural populations. Heterozygosity may well be maintained in populations by the action of heterosis, deleterious re- cessive genes, or even balanced lethals, especially in taxa such as C. berlandieri where translocation heterozygotes comprised 85% of all plants examined. In its high frequency of naturally-occurring translocations, Calylophus resembles some species of Gaura (Raven & Gregory, 1972b), Clarkia amoena (Lehm.) Nels. & Macbr. subsp. amoena (Håkansson, 1942), and some populations of Clarkia un- guiculata Lindl. ( Lewis, 1953) among the Onagraceae. Tetraploidy was observed sporadically in some subspecies of Calylophus hartwegii, but nowhere else in the genus. Seven populations of C. hartwegii subspp. hartwegii, pubescens, maccartii and intermediates among them were tetraploid (n = 14). These plants all showed a combination of bivalents and multivalents in meiosis. No taxon was found to be wholly tetraploid, and any evolutionary significance is probably minor. Tetraploid individuals appear to arise within populations from time to time. Extra, diminutive chromosomes were found in several taxa from both sec- tions of Calylophus, and could well occur in all of the taxa. They are small dark- staining bodies that often appear to be heteropycnotic and tend to proceed through first meiotic prophase and metaphase more rapidly than the normal chromosomes. Pairing between diminutives was common, and possible associa- tions with the larger chromosomes were seen occasionally. The most frequent ob- servation consisted of a single diminutive pair in addition to the normal comple- [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN eT jo Sup d + "T 6FI 9 jo Bult [ + "p ‘p JO suu Z + NE FSI (suore[ndod ez ) sniofiuid ‘dsqns n1421pupj42q 2) (N) euxo FRI uc:(X)*nxog-rFI-—Uuc'(5)rjosuug Tc £FF£c fi408247) Q zunJy TI J0 BUET + "I I6I 9 jo BUU I + "p 061 liiis e + 9 30 I-860S9 ee ? HDYog r jo sBuri tue M jo suu I + ue 'oO u[oour'q '07) 193sn7) euroqepriO 00 UOSTIAA 0 0 AORTA o) ?LIOJOLA 00 quioosdrq ‘og Apousy 00 IId 00 AopaeH ‘OFZ Lze (5) 8josuu [ + "c 9 Jo BUN [ + p jo Sun [ + "NZ ‘9 jo Bu [ + "p ‘(sued 8) 12 ZI jo Sung [ + "Tp jo sour 6 + g E (Sueſd g) p jo Bun [ + ''g u, FIFOI 1221011 "urup I + p jo Zur [ + Ug urup g + Uy 861 F JO sun g + NE 991 9 jo BuU [ + p jo Suri [ + "g Gol 95/9 8 jo BUN I + Jo sured, + UI SE sHəqoll p jo BuU T + 18 + "p F0&6[ fi408247) Q uaavy 9 Jo BULL [ + p jo BUN [ + "g ‘p jo sun T +g 64I 9 jo Bull [ + p JO sñuu g “surup Z€ + p jo ssuu ç + "NE 227 ‘surup € 9 JO ureqo I + "p I9 ty, e[qeqoid 09 sərrpiəumusrədns 70} I + 8 Jo BUN I + F JO Sun [ + "p p jo spun g + uf OFT (¿+ "q)] = u SET 9 J0 Bun p + "p Z8 ‘surup c + 9 jo BULL [ + F jo BUU [ + "gp JO Burd [ + MG i") Ic GSC Modan Q zunJq 081 (suone[ndod zz ) uaipuppiag ‘dsqns 1uuƏ1pupn]iaq 9 `oO unoq[eO 'oO S4oo1g sexə L 0D uui L euioqepro `oO Appa 00 vorg A OOITX9]N MƏN SuoneA19sq() uorpo[oo Aje „Snydojfjpo Jo suore[ndod ſvamqvu jo A30[0347) "T ATAV L TOWNER—CALYLOPHUS 53 1977] p jo Zulu + Ne IZI 00 9OURLIOT, F Jo Bunt [ + "Q 6II ‘OZ 0930105 F Jo soul c + NE ‘p Jo Bun [ + Mg 88 "091970 F jo gun g + NE FPG O PEE) (9) pjo Sut p + c O POBUD Q zun oO SAL OOIXO9]N MAN yd ureyo [ + p JO Sut [ + t€ “surup p 03 ë + F Jo Sur ng ELE ‘OD equoedy vuOZ AN (suore[ndod cT) uojpuof *dsqns ndomgnq['2 p Jo BUET HNT Hp 72 “OD OBL TIM, 9 Jo BUILT + up IZ 00 SIEN ‘surup 6 OFT + F JO SUN [ + 1e (Sued €) F Jo BUN [ + Uo) SCI HoBao Q uaavy ‘OZ peuo (59) 9 Jo BUU [ + F Jo BuU [ +g [EFES fi408247) Q zun OD uoo[[moojs CHa 9991 21X ‘0D [PPW (€ jo Suri T + 37 + y 10) e Jo Sun I + !T + ug GIF fi408242) 9 jo Bult [ + uc tg Jo Burr [ + "p 89861 fi408247) Q ua (T8149 Jeyonoa əsnouuəə:8 ) g jo 8un ¿c I + re Jaquinu ou ‘sowg 00 SKA (NA) F Jo sun 1 + uç 299 I %, 00 ardsso[ro) (X) F Jo Burt ng 06FS umbsopg "T oL bs e Lada! p jo ssull Z + e eJ 00 UMOIgG p JO Bult T + MG 49 F JO Sung + rr] 99 0 oourlg (M) F Jo ssun c + "€ PLOT WX (M) p jo sunt [ + "c SLOT Ut2]N (B) na [291 219 `9O 1exegq CI) 9 Jo nun p + rg 969I SINAT 'JN 2 `H oO douseg (XT) umo4g A "JN &[e»o[ ou Sexo] F Jo gun g + NE 6ST F jo soul g + iç ‘p Jo Burt ¿ rc ISI F jo Sur ¿ + UG c fi408242) Q uaavy 00 ueso SUOI]PA198Q() uoroo[on AY[ROO'T (penunguoo) I AL.. [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN 54 ‘surup ë + fF jo BULL [ + !!ç *àururp T + p jo BUU [ + "o ‘(sued g) 17 VSOSFI 220] paaig p J0 Bun [ + rg" 2769 ?ao]paaig y jo Bu [ + "ig “surup p + "4 88e "L OS p jo Bun [ + "e “surup Z + 17 67 "L GOSPI 2a0j;paaig ng + ug + Up “Se pp = u 08991 t[2nDA9]N ( suonepdod II) usanyoy `dsqns 3Bəm1mi `D sufunp [T 03 L + p JO suf g + NE “surup 6 + p JO gun [ + Uç “urup I + cT Jo Zuu [ + "p “surup € 03 [ + ML 69 Buta] p JO spuur 6 + ue 6SI6I fi408242) Q uaavy y jo sul T "G 761 1. (O) v jo sou c ug SEEES fi403247) Q zunjy p jo Bult ng c p JO soul c + NE 6I (9) g jo Bur [ + F jo BUU [ + "g ZSg86 ñiuopərD Q ZUNW L= u SEI (suonve[ndod g) snyofyy ‘dsqns n32mjgany Y ogue1n(T iin uu. sojuoreoseng y OOIX9]A ‘OD IUM O0 PIEM sexo 00 01910 0 Appa '07) saaryD OOIX9[N MIN (sued g) 117 pa surunp S 0} p + p JO BUN T + Ng p jo Sua [ ug 96 p jo 2un [ 18 5n, F96I ut uosyop[ O N "n, 0£-.L-89 1]2U40d 00 OIpIsaig Sexo, p jo Buu g 8 iu, 98 sufuip p + F Jo Bul [ + ''G cg H) I9 ¿ + Mg ‘9 Jo suu [ + !!F 62 00 13319 WOTRO p jo sBuu c + Ug ‘p Jo Buu [ + Ng EZI SuoneA1osq(Q uo Ay[e207] (penunuoo) I s'T8V I 55 CALYLOPHUS TOWNER 1977] (9) r jo Bun p + ug I cee HDH Q zunpy . ` (2) 90 Sun I + p jo Sur [ + “g C6SES OBALO Q zung 9 jo BUILT + p jo Buu [ + Yg ‘p JO Buu ¿ + Ug cs (3); =u SOSES HOονννẽu Q zunjy ae r6 b er g^ GI F JO spurt g + e SOI p Jo aur Hig I9I p Jo ssu g +uc t u ZG (suone[ndod gq ) suoosaqnd qu `O M@ 101 “so +I =u CF Zaad u^ sb 1125 ce (sjue[d Z) f Jo ssun c + re 99861 fit03247) Q uaany p jo Sufi + Uç FE (sued g) F jo sult [ + o “urup I + "y t; 666 12410415 NJ cs F Jo Burs [ + "g 5 88 98 MT ig + 8. pT =u STEET HD e l (sued g) F Jo sun g + ng turp I + y iu ee (suonep[ndod Of) 134022Dut *dsqus 13omginy Y p jo sun, + "c SSFSI 23o]p221g F jo Sun p + Mg in FFI ?ao]poo4g H =u SESFI a0] paarg (u) 9) FIFE SUAM suoesosq ; n UOHnoo[[07) 00 oyouop euroqepiO 00 Kun 0 soaryy OOIX9JA MON '07) ZNIQ ejueg ‘OFZ 9st207) €UOZLLV '0D) gedez 'OD) APIA IPA Sexo, sedi[neure L uoo'] oAonN uh,“ OOIX9]N SO IVO AYL]ROO'T (pənunuoo) '[ 41 gv AL GARDEN [Vor. 64 ` 4 ANNALS OF THE MISSOURI BOTANIC 56 (sued g) p jo uri [ g sy 62 00 10 89 SeXo[ p jo sunu Ing SEITE uosi3puy 000 S910[O(T Op*10[O7) 'sururp ë + !! L SII oO ofeaeN =u FII 07) ouruo52oo vuozuy (suone[nudod ) snyofynpupan Y (uss 1ot[onoA ou) (H) "4 Jaquimu ou 'zunjq 'oo 9p[e^n (qnd pue? ‘upy ^^19q) (J) flu I0F£c fozas Q zunJq 00 10 89e Sexo[ (fpf 9 `oopu, `Aqəq ) ua IT NOT “Se “PI = u 68 W OOIX9]AN (A X ‘osaqnd *^39q) (5) F Jo Buu [ +g Ig fi408247) Q zunJq 00 adnjepeny OOIX9JA MON (qnd pue? ‘upy *^39q) ZZ 0} 61 = UG ‘p Jo BUU + 8 Ë (qnd pu? ‘uoy *^32q) (J) L =u 640I S2] OD [etd euOZLIV (suonv[ndod g) n3anuvy Y snooue[[29St JN p jo Bull [ + "Q 99 OO I9[99QA, 8 jo Bun [ + "iç ‘p Jo Bun [ + Mg 5717 9 & fi408247) np 9899 umbsojy "T 2 `.1 00 Ile p jo Bull [ + 'c 66 00 191104 nn + Ap + 41g + NE “Be [I =u COFSS li40824*) Q zunyy 00 500094 p jo sñuu c + g fp Jo Buu [ + !!G LIZ6I fisodary Q uaany 07) UOLIT (9) 9 jo ureuo [ + "p F9££c ogar Q zunJq 00 uosiəq[no SuOHrA SO uonoəə[[oƏ Áe (penuguoo) I Aa. 57 JALYLOPHUS TOWNER—C ZT JO BUN o[quqoad + U I :FT jo Bult o[qeqoad FT Jo BUN o[qeqoad pp Jo ureqo — (M) FT JO BUN FT Jo Zum o[qeqoad FI jo sur FI Jo Butt o[qeqoad €I JO Sut o[qeqoad + u ‘pI jo Burt o[qeqoid FI JO Zum ə|qeqo.d FT JO BUU o[quqoad FI JO Burt o[qeqoad FI Jo suit o[quqoad FI Jo sult HI JO sult (sued Z) HI Jo Zum TI Jo ureyp o[qeqoad. + uy (sued Z) FI JO sur (sued Z) FI Jo Burt o[qeqoad FI Jo BUN o[qeqoad FT Jo BUN o[qeqoad FI Jo suit o[qeuqoad FT Jo SUU o[qeqoad (M) FT Jo sut — Surunp c + 9 jo Sur į + p JO Bunt [ + rz °F jo Suff i g ng 'suturp p + p JO Bun 9 Jo Sun T + p Jo Sun p + ug u SUOHAJOSQ() £666] li403247) Q uoany 09I€ unzingy Q uinbso]v E£SF6] lito3247) Q uaavy ZSI 00 SESUuUI V sexə L 00 ug NV quos 881 0 0 SIIIN N OFT SPI `o) ARLINY OSI '07) urg SFI ZFI 00 u[oour'T 88 OS 0 40019 881 00 oureg BULOYRLYO 881 cel 'O7) 3[249s00q 181 '07) BONG d FE 00 soaryy OO N MƏN 091 '07) HOIS d uosiopuy 00 ptu 681 00 ploy susury 6-69 ]/DYSID]Y I 11045140W POEN vprur2 4ppupng vo ON suone[ndod og) snjp]na2s 77) 268 uosiopuy OO 9utq OYM FOI 101 ‘OZ YARD PpPAdN uo AYL[RIO'T ( panumnuoo ) I Av [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN ( ) š ) *urg1iəoun A1nuəpi ‘uses uəuroəds 19qonoA ON q 2861 “unig puri = (g X T) ‘OSGI 'ue3eH = (H) 8861 “Te P STK = : “Te Jo ryseAeqein = (X) ‘peysyqndun usarAH = (W) ‘O96T “Uy X AIOBAIQ = (2) “Ssaxo[[oOj Se pojejouue sso[un umo Kur OIE ea Te Lo sli ud Se ed la su soie 1 5 3 (Suvſd p) u) OFG6I fisodaizyy Q uaavy `oo uojd[ (9) MZ 68988 %u ꝙ aun 00 ofpfS, p jo Buu [ ug 0808 vosiapuy '07) 10% Sexo], p jo Buu [ + "Q 8I p Jo Buu [ + uc ZI Ambos 9T p jo sun, + uç ST u FI (5 ‘sued c) p jo Buu [ + "c SS£88 “OSEET fiu03247) 2 zunjy ‘0D Appa OOIX9]JA MON (suone[ndod 6) pn]no1iqn1 dsqns njnoiqnj Y (syuejd g) p JO ssuu c + Ue 6II PdA spsupy fo `n uoo AN OOIXO9]N ( uore[ndod I ) snsonz144s `dsqns paqna * 2) 9 jo Bull [ + "p ‘p JO sduu c + NE 201 F JO ssun ç + g 90I "OD BPO) euozliy (suonv[ndod g) thawno} ‘5 (M) FT Jo Bur Sg upzijn jy 2 u¹j H 0 ipsius UIUIOÁ AA FT Jo ure: (Sued c) pp jo Bur FSI oO orougeq urg FI JO 8uu o[qrssod FLI 00 €p108 eje N ‘surup c + ZT jo Bull + m ZI OO uosyoef FT JO Zuni ə|qeqoid 981 ‘OD u?1q907) GI jo 2uu o[qeqoud + HI ‘fT JO Sun o[qeqoud 98I PI Jo suu 281 ‘OZ uow SUOHEAI9SQQ) uonoə[|o25 Ay[e2071 (penunuoo) I alavy 1977] TOWNER—CALYLOPHUS 59 ment. Numbers of extra, diminutive chromosomes ranged from 1 to 11, but the numbers were highly variable within populations and seemed to vary even among separate determinations from a single plant. The extra, diminutive chromosomes of Calylophus differed from those reported in Gaura (Gregory & Klein, 1960) and in Oenothera (Cleland, 1951, 1967; Cleland & Hyde, 1963) in often being heteropycnotic; pairing of the diminutive, extra chromosomes was frequent in all three genera of Onagraceae. The extra, diminutive chromosomes reported by Ostergren (1947) in Anthoxanthum were heteropycnotic like those in Calylophus. Cleland (1951, 1967; Cleland & Hyde, 1963) has hypothesized that the extra, diminutive chromosomes in Oenothera hookeri Torr. & A. Gray, may have been derived following hybridization between this species of Oenothera and an entity belonging to another group of the genus. This appears doubtful since it has not been demonstrated that the chromosomes of the different groups of Oenothera differ significantly in size, or that their differences are or would be maintained in hybrids. Certainly, it would be very difficult to construct an analogous hypothesis for similar chromosomes in Calylophus and Gaura. In Calylophus, supernumerary chromosomes of normal morphology were found only in one population of C. berlandieri subsp. berlandieri (Towner 140), in which plants had one to two extra chromosomes. These resembled super- numeraries as found in Clarkia (Lewis 1951; Hakansson, 1949), Camissonia (Raven, 1962), Gaura (Gregory & Klein, 1960; Raven & Gregory, 1972b), and Gayophytum (Lewis et al., 1958). Inversion differences were encountered only in some experimental hybrids between C. berlandieri subspp. berlandieri and pinifolius, and also in some crosses between C. hartwegii and C. lavandulifolius. No evidence of inversion heterozygosity in natural populations was obtained, but it could well occur as an infrequent event. Chromosomal determinations from experimental hybrids indicated that the taxa within sect. Salpingia have become differentiated by no more than 3 re- ciprocal translocations and, rarely, by an inversion. Among C. tubicula and all subspecies of C. hartwegii, crosses showed either complete homology or 1 to 2 translocation differences. Calylophus lavandulifolius differed from the above by 2 to 3 translocations and an inversion. The two subspecies of C. berlandieri, on the other hand, have become strongly differentiated, based on the current evidence. Hybrids between them were heterozygous for 2-6 translocations and sometimes for 1 inversion. Moreover, geographically separated populations of C. berlandieri subsp. berlandieri showed differences of the same magnitude. Reciprocal crosses of C. serrulatus and C. berlandieri produced hybrids with 3-6 translocation differences. Lastly, the few determinations from intersectional hybrids between C. tubicula and C. berlandieri demonstrated differences of at least 5 translocations. In the hybrid plants meiotic chromosome pairing was variable and poor, while anaphase movement was irregular. The high degree of sterility seen in these and other intersectional hybrids was probably derived from such chromosomal causes. In contrast, hybrid pollen fertility was moderate to high for most intrasectional crosses. Lower hybrid fertility in certain intra- [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN “pees poj»o[[Oo-p[eij uo UMOIS sjue[d uo peuuojied o1aM Ámrrqneduroour-j[os 10j sjs93 ‘sased JSOU UJ v OS FSI saunoy sexo] “OD oroujeq urs OS 281 42umo sexo ° ⁰ UOIO9UIPT) OS £6£6I fi408247) Q uaavy sexo[ “OD sesue1y OS GIFS uosi2puy BWOYRTYO “OD euoueuro7) sngp|n.4as `D IS CGI fi408242) Q uaavy sexo “oO peua IS 89861 fi408242) Q uaavy sexo], OO SÁEH snyofrud dsqns waipuppiaq 72 IS I-9660S9 d4ou L 2 4400uog sexo “op n A IS FIFOI 1120Y sexo, “OD quioosdrT I BEI os]əd sexo “oD Indoor ` I CE 5142004 sexo ^07) Khun uaipuppiag ‘dsqns u21pupj42q 0 IS 6II Pd spsupwN fo `ñ OOIX9]N e] OAƏnN snso]n81qqs "dsqus pjnaign} ' 7) IS OFGGI fii08247) Q uaavy sexo], OO uod 18 0808 uosi2puy sexo L “0N 10 9 pynargny `dsqns n[noiqnj 2) IS ZOI 42umo, BUOZILY “00 osrq207) „Hi IS 268 * BPBAAN “OD 9utq NYM snyofynpuvan] “9D IS ge fisodasy sexo L ^07) [[2..19 ], IS SON 5 zun sexa L *07) S009 IS IIGGI fi408242) Q uaany sexo] OO uou suəosəqnd 'dsqus ndoqgny “9D IS F961 ut uosqop[ sexo “OD pd " IS ZII 42umo[ vuozuv “op oqoedy uajpuof 'dsqus nBəmunti : O IS 69 Suq] Sexo [, “OD PNUM IS 6SI6[ fi408247) Q u sexə L “oD prem IS SSO ñiuoBar Q zuny OOIX9JA MAN “OD 0191O OO dsqus uBəm1im] 72) IS GF Zaad sexo L “oo ejedez IS 9886 403D Q uaany sexo “OD OPU ueg IS 66c 42410144S sexo, “or Aouury nyvosvu 'dsqns h “OD IS 09991 YanvAIW OƏINƏJN *soguor[eosens y IS SOS ?ao]paoig OxI *oSiuein(q IS SVS ?ao]paog OOIX9]N 'se29jvovZ IS FFEFI HO paas1g O9IX9]JA “seoə1eoez IS SO8FI ?ao]pooig XƏ *enuenum- nSomjny `dsqns usanppy `p sj[nsoy uonoəə|[o2 A[e2o7T uox? |, &ÁAj[rqueduroour-j[os 103 peururexo suone[ndod sn«dojfipp?)) c a18v L 1977] TOWNER—CALYLOPHUS 61 sectional crosses involving C. lavandulifolius and some involving C. berlandieri was correlated with cytological differences between the parental plants. FLORAL BIOLOGWYW AND POLLINATION Information on floral biology is based on the field work of D. P. Gregory (1964 and personal communication) and P. H. Raven (personal communication ), and on my own field work and study of cultivated plants. Collections of flower visiting insects were made, and these are to be deposited at the California Insect Survey, Berkeley. Determination of ultraviolet reflection and absorption pat- terns was carried out by photography with black and white film under near- ultraviolet illumination. All taxa of Calylophus were tested for self-incompati- bility by making repeated attempts at self-pollination. Table 2 lists those populations which were tested. The breeding systems of Calylophus are of three basic types. That of C. ser- rulatus is based on complex structural heterozygosity. In this species the flowers are highly autogamous, often self-pollinating before anthesis (Fig. 6). For this system insect visitation and pollen transfer are unnecessary, and are in fact un- common in C. serrulatus. The other two types of systems involve self-sterility and insect pollination. In sect. Salpingia, with the exception of C. tubicula, flowers are adapted for vespertine pollination by hawkmoths. They have narrow floral tubes measuring 25-50 mm in length (Figs. 1-3), sweet-scented nectar, large exserted stigmas, vespertine or afternoon anthesis, and, except for C. toumeyi, strongly ultraviolet- reflective areas on the distal portions of the petals (Figs. 7, 10). These reflective areas on the petals contrast markedly with small ultraviolet-absorptive regions which are usually present in the center of the flower. Populations of the tubular- flowered taxa experience vespertine and nocturnal visitation by hawkmoths, especially the abundant and effective pollen vector Hyles lineata (Fabr.) (Ce- lerio lineata). Some taxa and populations within taxa of C. hartwegii tend to have mid-afternoon anthesis and large- to moderate-sized ultraviolet-absorptive areas in the center of the flower. These seem to be secondary adaptations for bee pollination. Many populations of taxa in sect. Salpingia are visited in the afternoon, early evening, and even morning by halictid, anthophorid, and other bees. Some of these are oligolectic for the Onagraceae and probably contribute to pollination. The late-opening subspecies of C. hartwegii (Figs. 9, 10), along with C. lavandulifolius and C. toumeyi, have ultraviolet patterns less appropriate for bee pollination. The last species has no strongly ultraviolet-reflective areas on the petals (Figs. 11, 12). In these taxa bee visitation is restricted to the early eve- ning and is probably less effective in pollination. Calylophus tubicula and C. berlandieri are pollinated by matinal and diurnal insects. Their flowers open in the early morning, have short funnelform tubes ( Figs. 4, 5), and display large ultraviolet-absorptive regions in their centers ( Figs. 13, 14). The petals are highly ultraviolet-reflective distally. Based on the available evidence, the primary insect visitors to C. tubicula seem to be morning-active halictid bees. Other potential pollinators include hawkmoths, which visit the flowers lightly at about sunrise. 62 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 < Ficung 1. Calylophus toumeyi in the Chiricahua Mountains, Cochise Co., Arizona (Towner 107). A great variety of insects come to the flowers of C. berlandieri, which appears to possess a generalized pollination system. Beetles, skippers, small butterflies, occasional hawkmoths, and a wide variety of bees have been observed gathering pollen or nectar. Each of these groups may contribute to pollination, with its relative importance varying with locality. When present, ultraviolet patterns on the flowers serve to direct insects to the anthers and nectar. The pattern typical of Calylophus consists of absorptive regions at the base of the petals, at the mouth of the floral tube, and on the stigma and anthers (Figs. 8, 10, 14). Calylophus toumeyi differs in having moderate ultraviolet absorption over the entire flower (Fig. 12). In Onagraceae, ultra- violet absorption results from the presence of one or more flavonoids with ab- sorption maxima in the near-ultraviolet (Dement & Raven, 1973, 1974). As in many other yellow-flowered Onagraceae, the ultraviolet-absorptive portions of the petals of Calylophus hartwegii and C. serrulatus, and presumably those of the other species as well, contain isosalipurposide, a chalcone with an absorption maximum at 365 m, (Dement & Raven, 1973, 1974, and personal com- munication). In addition, C. hartwegii has an accompanying flavonol, myrecetin- 3-glucoside or galactoside, and C. serrulatus has an unidentified compound that resembles an aurone, a class of flavonoid that is frequently associated with chal- cones in Asteraceae (W. Dement, personal communication). of these flavonoids are absent from the ultraviolet-reflective portions of the petals. Carot- enoids absorbing maximally at 400-470 m, are found throughout the petals. 1977 TOWNER—CALYLOPHUS 63 s ii. —3. C. toume vi ie . tubicula subsp. tubicula.—5. C. berlandieri subsp. pinifolius. 3. C serrulatus. All x Ficures 2-6. Longitudinal sections of flowers of Calylophus.—2. C. hartwegii subsp. C Therefore, insects with trichromic vision would see the petal apex color as “bee yellow” and the petal base, anthers, and stigma as “bee purple.” It is uncertain whether these ultraviolet patterns would be as effective for hawkmoth pollination after sunset as they are for diurnal insects in sunlight. The small size or absence of ultraviolet absorbing areas in the evening-opening Calylophus perhaps signifies that such pattens are not useful for insect orienta- tion at night. Alternatively the minimization of ultraviolet contrast patterns may have evolved to lessen pollen and nectar removal by diurnal insects inefficient in pollinating the vespertine Calylophus. This could be especially true in the 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 URES 7-12. dpi is Calylophus under fluorescent and ultraviolet illumination. Collections refer to seed sources.—7. C. hartwegii subsp. hartwegii (Durango, Mexico, Breedlove 14305A); fluorescent illumin: d emasculated.—8. Same, with ultraviolet illumination.—9. C. hartwegii subsp. fendleri ( Presidio Co., Texas, jatari in 1964); fluorescent illumination.— ` . Same, with ultraviolet cu i ımeyi (Cochise Co., Arizona, Towner 107); fluorescent illumination.—12. Same, with 1 illumination. case of C. toumeyi, which has flowers contrasting only slightly with its vegetative parts and the background in the ultraviolet region of the spectrum. Mazokhin- Porshnyakov (1969) states that considerable ultraviolet light is present at night, especially in moonlight. However, the moths investigated have shown low sensi- tivities to ultraviolet light, and become functionally colorblind at light levels below 0.05 lux. Their maximum sensitivity to light is in the vellow to green 19771 TOW NER—CALYLOPHUS 65 Ficures 13-14. Flowers of Calylophus berlandieri subsp. berlandieri (Hartley Co., Texas, Roberts 35) under fluorescent and ultraviolet illumination.—13. Fluorescent illumination.— 14. Ultraviolet illumination. region of the spectrum. If hawkmoths follow this pattern, they could locate Calylophus flowers merely by their scent and reflectivity at wavelengths greater than 480 my, eliminating any need for ultraviolet reflection. With the moderate light levels present at dusk both bees and moths could probably utilize the ultra- violet contrast patterns for orientation. PHYLOGENETIC RELATIONSHIPS The species of Calylophus fall into two clearly recognizable but related groups. Those with keeled sepals and two sets of stamens of unequal length are included in sect. Calylophus and those with plane sepals and subequal stamens are as- signed to sect. Salpingia. No intermediates between these sections have been found, and the species of the two groups are intersterile while being relatively infertile among themselves. Fifty-one of 101 intersectional crosses yielded some seed, with the amount set ranging from 5 to 81% of normal, but germination was poor ( 1-3% ) and the few weak hybrids that grew to maturity were largely sterile (0-10% pollen fertility). Members of sect. Salpingia are most likely primitive in comparison with those of sect. Calylophus, as they have a clumped perennial habit and, except for C. tubicula, are primarily pollinated by hawkmoths. In growth form, the members of sect. Salpingia resemble the more generalized forms of Gaura (Raven & Gregory, 1972a), a genus which is also primarily moth- pollinated and closely related to Calylophus. Among the hawkmoth-pollinated members of sect. Salpingia, C. lavanduli- folius seems to be the most distinct, having a more caespitose habit than the other forms and a broad geographical distribution which includes numerous relict populations. Calylophus toumeyi appears to be most closely related to the southern subspecies of C. hartwegii, especially C. hartwegii subsp. hartwegii, with which it shares long free sepal tips, preference for montane habitats, and strigulose pubescence. Within C. hartwegii a pattern of reticulate relationships is evident and relative affinities are difficult to assess. Calylophus hartwegii subsp. maccartii, occupying the Texas coastal plain, is likely a recent derivative of 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 C. hartwegii subsp. hartwegii, as these two forms share several characters and are geographically adjacent. Calylophus hartwegii subsp. pubescens, C. hart wegii subsp. fendleri, and C. hartwegii subsp. filifolius seem to constitute a series of related forms occupying slightly different ecological zones in the plains of the southwestern United States. The subspecies of C. hartwegii are connected by zones of intergradation, although not all of the possibilities for geographical con- tact and gene exchange are realized. The short-lived perennial C. tubicula is unique in sect. Salpingia, having several characteristics of bee-pollinated flowers. It is likely a specialized derivative of relatively recent origin. Sect. Calylophus is apparently more specialized in having an annual or short- lived perennial habit, and perhaps also in its adaptations to diurnal pollination. Of the two species in the section, C. berlandieri is clearly the more generalized, having a self-incompatibility system, and chromosomes which form bivalents or small rings at meiotic metaphase I. Calylophus serrulatus, a highly autogamous complex structural heterozygote, is a specialized offshoot of either the former species or a common ancestor. No clear knowledge of the exact ancestry of C. serrulatus is available, although hypothetical lineages have been worked out for some of the other complex heterozygotes in the Onagraceae. It may be that this species developed through the independent origin of complex structural heterozy- gosity in different geographic regions. This is suggested by the fact that neighbor- ing populations of C. serrulatus and C. berlandieri often tend to resemble one another phenetically, a condition which could be attributable to multiple an- cestry for C. serrulatus, to introgressive genetic exchange, or to parallel responses by the two species to local selective regimes. Calylophus serrulatus is highly suc- cessful and widespread in the Great Plains from Canada to New Mexico and also occurs in Arizona and Mexico, Calylophus berlandieri is largely restricted to Texas. A likely but speculative evolutionary sequence could have begun with the primary divergence of the ancestors of the two sections. Accompanying. this split was the development of morning anthesis and other characteristics of day- pollinated flowers in sect. Calylophus. The moth-pollinated perennial ancestors of sect. Salpingia probably evolved more slowly, but gave rise to C. tubicula. Calylophus serrulatus presumably developed very recently from C. berlandieri or its ancestors. The course of evolution within sect. Salpingia seems to have depended upon ecogeographical differentiation among the various forms, while being little dependent on cytological transformations. Cyclic climatic fluctuations in Pleistocene and Recent times have undoubtedly contributed to the taxonomic differentiation within sect. Salpingia, judging from the confusing patterns of over- lapping geographical pas isolated populations, and introgression that are now in evidence among its tax Divergence within sect. C nae has been marked by much greater cyto- logical change than in sect. Salpingia. Several chromosomal rearrangements involving translocations were required for the derivation of C. serrulatus from pair-forming outcrossers. Similar chromosomal repatterning has also occurred within C. berlandieri, and characterizes at least three series of populations. In addition, the two races of C. berlandieri appear to be ecologically differentiated, 1977] TOWNER—CALYLOPHUS 67 and remain well separated in regions where there exists a discontinuity between their respective habitats. TAXONOMY Calylophus Spach, Hist. Nat. Vég. Phan. 4: 349. 1835. “Calylophis” "us Nouv. Ann. Mus. Hist. Nat. 4: 337. Nat 35. Meriolix Raf. ex Endl., Gen. Pl. 1190. June 1840: Raf., Amer. Monthly Mag. & Crit. 192. 1819, nom. nud.; Raf. J. Phys. Chim. Hist > Rev. 4: . Nat. Arts 89: 259. 1819, nom. € Meriolix emra lutu (Nutt.) Walp. — 5 serrulatus (Nutt.) Raven. Salpingia (Torr. & A. Gray) Raim. in Engler & Pri antl, url. Vy n 3(7): 217. 1893. Based on Oenothera subg. pores Torr. & A. Gray, n N 1: 50 O. TYPE: J. Oenothera lavandulaefolia Torr. A. Gray = tn TE ee (Torr. & A. Gray) ; Galpinsia PEE: 5 7 Torrey Bot. Club 5: 236. 1894. Based on Oenothera subg. Salpingia Tor A. Gray, Fl. Amer. 1: 501. 1840. rype: Galpinsia hartwegii ( Benth.) Britton = E a i oon ( Benth.) Raven. Herbaceous to suffrutescent perennials, rarely annual, from a woody caudex, flowering in the first year. Stems nearly prostrate or decumbent to erect, with grey to pinkish brown epidermis, this sometimes exfoliating. No basal rosette, leaves cauline, more or less sessile, alternate, entire to spinuose-serrate, the up- per leaves more or less uniform in size, the lowermost often somewhat larger: stipules absent. Flower 4-merous, actinomorphic, borne in the axils of the upper leaves, opening in the early morning or from midafternoon to near sunset, with the stigma receptive and anthers shedding pollen simultaneously upon anthesis or soon afterwards, wilting in 1'2 to 2 days; buds erect in inflorescence. Floral tube well developed and prolonged beyond the ovary, deciduous after anthesis. Sepals greenish yellow, often with purple or red markings, reflexed separately. Petals yellow, in some species becoming red, orange, or purple upon wilting, reflexed in anthesis. Style yellow; stigma yellow to yellow green, occasionally blue black in one species, peltate, discoid to nearly square, sometimes obscurely and shallowly 4-lobed. Stamens 8, yellow; anthers narrowly elliptic to linear, versatile, the sporogenous tissue divided into packets within each locule; pollen yellow, shed singly. Capsule many seeded, sessile, cylindrical, and often nar- rowed at each end, obtusely 4-angled, longitudinally dehiscent, persisting on the stem after dehiscence. Seeds in 2 rows in each of the 4 locules. Basic chromosome number, x = 7. Five of the six species are self-incompatible. TYPE species: Calylophus nuttallii Spach = C. serrulatus ( Nutt.) Raven. In the accounts which follow, the taxa will be grouped according to their phenetic affinities. Specimens cited were selected to represent the ranges of morphological variation and geographical occurrence. Where possible, I gave preference to recent collections and those with numbers of duplicates. My own collections are deposited in the Dudley Herbarium of Stanford University ( DS) with duplicates to be distributed. KEY TO SECTIONS à. Sepals plane, lacking a keeled midrib; stamens subequal section I. Salpingia aa. Sepals with conspicuously keeled m d; stamens biseriate, the epise palo filaments about twice as long as the epipetalous filaments ction II. Calylophus ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 68 KEY TO SPECIES Section I. Salpingia a. Floral tube O. in 9 75 upper two-thirds or more, or less than 15 mm as flowers opening near sunrise _ 4, tubicula aa. Floral tube er els in upper half or less, 15-55 mm long; flowers 3 near sunset. » With conspicuous axillary n of small leaves, these up to 30 mm long; subulate sepal tips 2-9( -12 n long; capsule thin walled and dehiscent only in the distal portion; montane porte northwestern Mexico to southeastern C. toumeyi = Arizona _ — 3 IRURE 3. bb. With or without axillary fascicles of leaves or if present, these less than 20 mm long; subulate sepal tips 0.5-6 mm long; capsule thicker walled, dehiscent along its entire length. Plants low, frequently caespitose, mostly 0.4-2 dm high; Ng: gra strigulose; sepal tips short, 0.3-3 mm long avandulifolius ce. Plants not caespitose, mostly taller and more openly mnt 044 —4 dm high; variously pubescent or glabrous; if "as sad is 2-6 mm ng mes . C. hartwegii ection A Calylophus a. Flowers small, the petals mostly 5-12 mm long; stigma positioned near the apex of the floral tube or slightly beyond, within the circle of anthers; 30-80% of pollen grains aborted irc edd ee a ae serrulatus aa. Flowers usually larger, the petals mostly 9-25 mm long; stigma well exserted. usually to the end of the 8 anthers or isi 85-100 mes of FEM grains fertile 5. C. berlandieri Section I. Salpingia (Torr. & A. Gray) Towner, comb. nov. ip ncs haat Salpingia Torr. & A. Gray, Fl. N. Amer. 1: 501. sagt e ( Torr. & Raim. in Engler & Prantl, Natürl. Pflanzenfam. 3(7): 217. 1893. Galpinsia 5 85 Mem. Torrey Bot. Club 5: 236. 894. Herbaceous to suffrutescent perennials 0.4-6 dm high, glabrous to glandular- pubescent, strigulose, or with spreading trichomes. Leaves 0.3-5 cm long, entire to serrulate. Inflorescence sparse to dense; buds terete. Flowers opening in af- ternoon, evening, or morning. Floral tube terete, tubular and gradually expanded through its entire length, or tubular proximally and funnelform distally, 5-70 mm long. Sepals plane. Petals suborbicular to rhomboidal or squarish. Stamens nearly equal in length. Capsule promptly dehiscent upon drying, usually not curved. TYPE SPECIES: Calylophus lavandulifolius (Torr. & A. Gray) Raven. 1. Calylophus hartwegii (Benth.) Raven, Brittonia 16: 286. 1964. Oenothera hartwegi Benth., Pl. Hartw. 5. 1839. Herbaceous to suffrutescent perennial arising from a woody caudex; stems one to many, sparingly to densely branched above, nearly prostrate to erect, 0.4—5 dm high, strigulose, glandular-pubescent, glabrous, or with spreading hairs, more densely pubescent above. Leaves sparsely to densely distributed along the stem, sessile or indistinctly petiolate, spreading to ascending, sometimes reflexed, linear or filiform to ovate or oblanceolate, 3-50 mm long, 0.4-12 mm wide, usually not much reduced up to the stem, variously pubescent or glabrous, the tip acute 1977] TOWNER—CALYLOPHUS 69 to obtuse, the base acute-attenuate to truncate-clasping, the margin entire to ser- rate, frequently undulate; fascicles of small leaves 1-15 mm long sometimes present in the nonfloriferous axils; lowest stem leaves sometimes wider than above, frequently obovate to spatulate, to 65 mm long. Inflorescence lax, with rarely more than one flower at a time fresh on a stem, variously pubescent or glabrous; buds terete. Floral tube terete, tubular in the lower one-half or more, gradually expanded distally, 16-50(-60) mm long, 4-20 mm wide at the throat of pressed specimens, variously pubescent or glabrous without, the inner surface glabrous distally, sometimes minutely pubescent at the base, frequently fading to pink or purple on wilting. Sepals 7-28 mm long, 2-10 mm wide, with subulate free tips 0.5-6 mm long, plane, variously pubescent or glabrous, pale yellow- green, frequently with purple spotting or marginal stripes, fading as with the floral tube. Petals suborbicular to rhomboidal, 10-35 mm long, similar in width, highly ultraviolet-reflective, with basal ultraviolet-absorptive spot of varying size, sometimes absent, frequently turning pinkish or purplish upon wilting. Stamens subequal; filaments 4-13 mm long, glabrous; anthers 5-13 mm long. Style 25-65 (-75) mm long, usually exceeding the stamens, minutely pubescent below; stigma flat to slightly revolute, squarish, 1.5-6 mm broad; ovary 4-30 mm long, 1-3 mm wide, variously pubescent or glabrous. Capsule 640 mm long, 2-4 mm wide, moderately thick-walled, completely dehiscent, straight or slightly curved; seeds 1-2.5 mm long, obovoid, rounded or sharply angled, truncate at the apex. Self-incompatible. Gametic chromosome numbers, n = 7, 14 TYPE: MEXICO. AGUASCALIENTES: Aguascalientes, 1837, Theodor Hartweg 10 (K, holotype; P, isotype). Distribution: Local and colonial to abundant and widespread on rocky, sandy, gypsum, or limestone soils in arid to relatively mesic open areas, in south- eastern Colorado, southwestern Kansas, western Oklahoma, Texas (except east- ern part), New Mexico, southeastern and east-central Arizona, and in Mexico from Chihuahua, northern Coahuila, and northwestern Tamaulipas south to Aguascalientes. From ca. 30 to ca. 2,500 m elevation. Flowers February to Oc- tober. As treated here, Calylophus hartwegii includes five intergrading subspecies. The species is distributed over much of the Southwest and northern Mexico, oc- cupying relatively dry plains and mountain regions. Long, slender floral tubes and vespertine anthesis characterize all subspecies of C. hartwegii, suggesting a basic adaptation to hawkmoth pollination. Varia- tion among the subspecies in exact time of anthesis, in ultraviolet reflection pat- terns, and in flower size was observed, and is likely due to modal differences in pollination systems. Principal flower visitors included halictid and anthophorid bees and hawkmoths, but with differing proportions of these insects visiting the various populations observed. The mean length of the floral tube in this species and in C. lavandulifolius and C. toumeyi is about 30-40 mm. Anthers and stigma are well exserted beyond the tube, but tend to block the entrance to it. Short-tongued hawkmoths, such as the abundant species Hyles lineata 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 (Celerio lineata), land on the flower and extend their heads into the tube to obtain nectar. In doing this they pick up quantities of pollen on the head and thorax and can serve as effective agents of pollination. All subspecies of C. hartwegii have been tested at least once for self-incompatibility. In a brief check, each of 22 plants examined was found to be self-sterile. Of 88 diploid plants from 55 populations examined for chromosome configu- ration, 56 individuals from 42 populations, including some from each subspecies, showed translocation heterozygosity. The mean number of heterozygosities per plant was 1.0 for the species as a whole. Plants with as many as 5 translocation heterozygosities were found in seed obtained from natural populations, but most individuals had only 1 or 2, displaying 1 or 2 rings of 4 chromosomes or a ring of 6 at meiotic metaphase I. Calylophus hartwegii is a predominantly diploid species, although occasional tetraploids and plants with extra chromosomes were found. Sixteen plants from nine populations, including all subspecies but C. hartwegii subsp. pubescens, had extra diminutive chromosomes. Five of the 62 populations from which chromo- some number determinations have been made contained tetraploid individuals. No population was found to be comprised of both tetraploid and diploid plants, although the low number of duplicate counts from tetraploid populations leaves this possibility open. One plant intermediate between subspp. pubescens and hartwegii (Towner 3) appeared to be triploid, further increasing this likelihood. Tetraploids were found in C. hartwegii subspp. pubescens, hartwegii, and mac- cartii, all predominantly diploid taxa. Difficulties of interpreting the variation in this species have arisen from a number of causes. The entities in sect. Salpingia tend to have extensive geo- graphical ranges which overlap and interdigitate. Minor ecological differences often separate the taxa locally. The subspecies of C. hartwegii are distinguished morphologically from one another by a few rather slight differences such as pubescence and leaf shape. The pattern of variation within the section is reticu- late, with few characters varying concordantly. Broad areas of introgression exist between certain of the taxa, while others show greater discontinuities in areas of contact. All forms retained in C. hartwegii are connected either directly or indirectly by intergradation. The center of the distribution of this species in western Texas and northern Mexico is characterized by an abundance of forms occurring in near proximity and by a complex display of variation. Absence of any significant number of intermediates in the appropriate regions indicates that C. hartwegii, C. lavandulifolius, and C. toumeyi should be dis- tinguished specifically. The earlier decision to combine them (Towner & Raven, 1970) was intended to emphasize the overall unity of this group, but a subsequent thorough study of herbarium material failed to bring forth evidence of intergrada- tion among the three species. Populations of C. hartwegii, as mentioned above, occasionally occur sympatri- cally with C. berlandieri, C. serrulatus, and C. tubicula. They also frequently occur mixed with C. lavandulifolius, and only rarely form hybrids with that species. 1977] TOWNER—CALYLOPHUS 71 KEY TO SUBSPECIES a. Ovary (and usually stems and leaf margins) with spreading trichomes; leaves (except lowest) abruptly narrowed to truncate or slightly clasping at the base; widespreac SNS S CENTUM. ~ RRM V Nn cabs sp. pubescens aa. Plant w vithout spreading trichomes: leaves gradually narrowed at the base or extremely narrow 1 out. b. Plan 4. a or nearly so; flowers opening from one hour before to one hour afte sunset; widespread FEE ld. subsp. m vll Plant, Panas on ovary and t upper stems, with some form of pubescence; flower usually opening 2-5 hours before sunset, occasionally later €; vary and stems with short glandular pubescence; leaves glabrous t glandular-pubescent, rarely sparsely strigulose, filiform to narrowly 1 late; gypsum or limestone flats, central New Mexico to northern Mexic CC le. subst » filiis y c - eC ó 2 ~ E ie [^7] 2 ° — — g e z — Qu — e e < G mn — n I — — < z; oe 2 af 0g x 8. = ° n © e ° = ii — ga E e Ë z = [o» 2 z — > T "3 [zi = ° uw [e] 6 = Pf the 15 ives narrow ly e olata to amelie or did Bis. d. Leaves mostly 4.5 to 11 times as long as wide, usually with crinkled- undulate margins; plant sparsely strigulose or 3 minutely S cea pubescent; low plains from southeastern Texas to northern Mexico _ — 1b. subsp. maccartii "p da mostly 9 to 35 times as long as ; wide, usui ally 1 not cı rinkled; plant sparsely to densely strigulose; high plains and mountains from southern rans-Pecos Texas to Aguascalientes la. subsp. hartwegii dd. Qu la. Calylophus hartwegii ( Benth.) Raven subsp. hartwegii; Towner in Correll and Johnston, Man. Vasc. Pl. Texas 1121. 1970.—Fic. 2 Salpingia hartwegii (Benth. ) Raim. in Engler & Prantl., Natürl. Pflanzenfam. 3(7): 217. 1893. Galpinsia hartwegii ( Benth.) Britton, Mem. Torrey Bot. Club 5: 236. 1894. Oenothera hae Benth. var. typica Munz, Amer. J. Bot. 16: 706. 1929, pro parte. Calylophus hartwegii var. hartwegii; Shinners, Sida 1: 342. 1964, pro parte. Oenothera hartwegii ar, hartwegii; Munz, N. Amer. Fl., ser. 2, 5: 139. 1965, pro parte. Oenothera greggii A. Gray var. pringlei Munz, Amer. J. Bot. 16: 711. 1929, pro parte. O. Uu ei (Munz) Munz, N. Amer. Fl., ser. 2, 5: 138. 1965, pro parte. TYPE: Bachimba n, Chihuahua, Mexico, 27 March 1885, ve G. Pringle 224 (GH). PURI win kondu i ola T orr. & A. Gray var. typica Munz sensu Munz, Amer. J. Bot. 16: 704 29, pro parte. O. lavandulifolia var. 55 e sensu Munz, N. Amer. Fl., ser. 2, 5: 138. 1965, pro parte. Stems several to many, sparingly branched above, decumbent to somewhat ascending, or plant tufted, 0.5-3 dm high; plant grayish-strigulose throughout more densely so on the ovary and inflorescence than elsewhere. Leaves dense on stems, more or less ascending, linear to narrowly lanceolate, 10-35 mm long, mm wide, the tip acute, the base acute-attenuate, the margin entire to shallowly and sparsely serrulate, occasionally undulate; fascicles of small leaves 2-15 mm long usually present in the axils. Floral tube (18—)30-50(-60) mm long, 4-13 mm wide at the throat, sometimes with purple longitudinal bands, often fading purplish. Sepals 8-20 mm long, 3-7 mm wide, with free tips (1-)2-6 mm long, frequently with purple marginal stripes. Petals squarish or rhomboidal, 13-30 mm long, frequently fading to a purple or pink color, with basal ultra- violet-absorptive spot absent or present and of small to moderate size. Filaments 5-10 mm long; anthers 5-9 mm long. Style 30-65(-75) mm long, glabrous: stigma 2-5 mm broad; ovary 5-12 mm long, 1-2 mm wide. Capsule 10-25 mm long, 2-4 mm wide; seeds 1-2.5 mm long. Self-incompatible. Gametic chromo- some numbers, n — 7, 14. 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 * PES | | l | | T =o | | | | | | | | | ° ° e . N 7 as ° | "he e. a ^ à ( ee = Pu vu Y ° °° 0 J des ^ c: yo * i e° Pe ae K x< 4 \ | k JS * e e % 2 ^ J ° ° N 8 4 ° ) e â L : e" ñ ç P4 ° RE P I * * N ° o VA .. Lf Sn „„ ure 15. Distributions of Calylophus hartwegii subsp. hartwegii (dots), C. hartwegii subsp. maccartii (triangles), and intergrades between the two subspecies [open circles). 1977] TOWNER-—CALYLOPHUS 73 Distribution (Fig. 15): Mostly on rocky or gravelly soil, sometimes lime- stone, in rugged canyons in the northern part of the range, to high plains and mountains, reaching pine forest at the southern limits of the range, from Brewster and southern Hudspeth cos., Texas, south through central Chihuahua, Coahuila, western Nuevo León, and eastern Durango to central and southeastern Zacatecas, Aguascalientes, and southwestern San Luis Potosi. Elevational distribution from ca. 900 m (near Solis, Brewster Co., Texas) to at least 2,300 m (6 mi N of Zacate- cas, Zacatecas, Mexico). Flowers February to October. 5 L: esas . STATE XAS: r Co. Eos just N of Mariscal Canyon, Big Bend 5 Park. [ws dr Correll. 24568 ( d ead of Fresn o Canyon, Big Bend Ranch, rell & Rollins 23672 (LL). Near Marathon, p oung 174 (MO, TEX). Hudspeth Co.: 1 Hill-Fox Hill area of the central Malone Mts., Waterfall 5830 (GH, NY). Val Verde Co.(?): Pecos R., Thurber 123 (NY). County unknown: Coyote Mt., West Texas, Havard die^ E CHIHUAHUA: 12 mi W of General Trias, Breedlove 15741 (DS). 5 mi S of E. “del Parral, [ow 15745 (DS). 13 mi E of Hidalgo del Parral, Breedlove 14305 (DS). Santa Eulalia Mts., Rose 11693 (US). Santa Eulalia Hills, Wilkinson 4601 NY). Ca. 30 mi NW of Chihuahua, Lee 59 (F, TEX). Vicinity of Chihuahua, 1 59 (F, MO, DI NY, US). Alberto, SE of Chihuahua, Pennell 18627 (Ny. PH, US). mi N of Parral, Waterfall 12514 (OKLA). Chihuahua, Le Sueur E (F, UC). cn Parral and Villa Ocampo, Weber d» Charette 11710 (car O). Gallego Springs (between Carrizal and Chihuahua), Wislizenius in 1846 (MO). Near Chikushos dien, in 1886 (F, MO, NY, US). 13 mi N of Ciudad Chihuahua, Breedlove 15736 (DS). 24 mi W EN 1 S of Chihuahua, Stuessy 1020 (TEX). 23 mi W of Chihuahua, aaa 247 0 (ARIZ). Near Chihuahua, Le Sueur 811 = 58 (F, MO, SMU, TEX, US). COAHUILA: vu s Palm Canyon, Marsh 1002 (F, OKLA, TEX). Mízquiz Marsh 1134 (F, OKLA, SMU, TEX). Cerro de los Arboles, Jermy ee S). 11 km NE of Jimulco, Stanford et al. 9 (ARIZ, DS, MO, NY, UC, WTU). m NW of Fraile, Stanford et al. 402 (ARIZ, DS, MO, NY, WTU). Siena Mojada Mts., Jones po role POM, US). Sierra Mojada, Jones in 1891 (POM). Near qom (now (un. eral Cepeda), Gregg 723 (MO). Near Buena Vista, SW of Saltillo, Gregg 387 (MO). Ciénega Grande (just NE of Parras), Gregg 492 1 5 b Alle nde, Marsh 1776 (F, TEX). Sierra de la Paila, Purpus 4977 (F, US). Sierra de Parras, Purpus in 1905 (UC). Saltillo, Arséne 6510 (US). Paso del Diamante, near Saltillo, Mis: 15034 (MO, POM). 30 mi W of Saltillo, Wislizenius 298 (MO). Saltillo (? » Palmer 344 (US). Saltillo, Palmer 337 (US). 9 mi S of Saltillo, Straw & Forman 1337 (RSA). Ca. = mi E of Saltillo, McVaugh 12301 (RSA). Saltillo, Fisher 32 (F, UC). Nuevo LEÓN: 12 mi N of Sabinas Hidalgo, Heard & Barkley 14535 (TEX). 4 mi S of “asw sap McGregor 5 a 65 (DS, KANU, SMU). punANcGOo: 46 mi b of La Zarca, Straw & Forman 1525 (RSA). 15 km NE of Guadalupe Victoria, Henrickson S). 77 mi S of Parral, “Wiens 3464 (COLO, DS). 6 mi NE of Hidalgo del Parral, Breed: 5947 (DS). 21 mi N of La Zarca, Mee 14305A (DS). 71 mi NE of Durango, all 133368 (OKLA, RSA, SMU). 3-6 mi W of La Zarca, Straw & Forman 1717 Men UC), Ca. 54 mi S of Villa Matamoros, Class Straw & Forman 2058 (RSA, UC). ZACATECAS: On route 49, 2 km N of Route 45, Cruden 1238 (DS). Near Concepción del Oro, Palmer 271 (F, MO, NY, UC, US). Gypsum flats, Sierra Hermosa, Shreve 8594 (ARIZ). i SE of Sombreto (Sombrerete?), Waterfall 13799 (OKLA, RSA, SMU). 7 mi S of Fresnillo Breedlove 15485 (DS). 9 mi N of Fresnillo, Breedlove 15486 (DS). 2 mi SW DS sn cepción del Oro, Pennell 17416 (PH). “Gravelly soil,” Purpus in 1903 (UC). Ca. 22 mi NE of 3 Straw & Forman 1492 (RSA). 55 mi W of Zacatecas, Re Ae et al. 2660 (DS). cm NE of Laguna Seca, on km 20 of road from San Luis Potosi to Antiguo Morelos Rzedowski 6325A (RSA). Charcas, Whiting 898 (ARIZ). Ca. 5 km ] ard je tehuala, ae 9186 (DS). 44 mi NW of San Luis Potosí on road 2 pee md 5954 (D qi 13 mi NW of San Luis Potosí on road to Zacatecas iu 143 27 mi NW of San Luis Potosí along road to 5 l 15473 (DS). Charcas, Landel 755 (ARIZ. DS, F, POM, UC, US). AGUASCALIE E of Aguascalientes, McVaugh 16680 (RSA, TEX). JAusco(?): Lake Chapala ifta almost certainly in error) Lemmon & Lemmon in 1905 (UC). 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Calylophus hartwegii subsp. hartwegii occupies a diversity of habitats, in- cluding desert scrub, thorn scrub, and pine forest. Typical material for this sub- species comes primarily from montane areas of north-central Mexico, while col- lections from lower altitudes and more northern localities exhibit evidence of hybridization with other taxa. Poor sampling in the remoter areas of northern Mexico has left details of the geographical range unclear. For example, the oc- currence of this subspecies throughout most of northern Coahuila and eastern Chihuahua seems likely, but is not yet established. The limits of the variation in this subspecies do not correspond closely to those set by Munz (1929) for Oenothera hartwegii var. typica. The strigose pubescence of Hartweg's type actually excludes it from Munz's description, and the bulk of the present taxon would also not be included. Oenothera greggii var. pringlei was named to accommodate the strigose-canescent forms, but since Hartweg's original type represents that group of populations var. pringlei must be reduced to synonomy. On the basis of leaf width, I have placed some elements of Munz's var. typica in C. hartwegii subsp. maccartii. In addition, a portion of Munz’s var. typica which included glandular-pubescent individuals is joined with C. hartwegii subsp. filifolius, as is part of Shinner's C. hartwegii var. hart- wegii. Lastly, some collections with glabrous leaves and stems placed in var. typica by Munz and in var. hartwegii by Shinners clearly belong with C. hart- wegii subsp. fendleri as it is here constituted. In all previous treatments, the composition of var. hartwegii has been extremely heterogeneous. Clear identifi- cation of the nature of Hartweg's type and the recognition of larger geographical assemblages have rendered the variation pattern for C. hartwegii less confusing, especially with regard to subsp. hartwegii. Finally, a few plants assigned to Oenothera lavandulaefolia by Munz clearly belong with C. hartwegii subsp. hartwegii on the basis of their long sepal tips, narrow leaves, and southern dis- tribution. Considerable variation in leaf width, stature, and degree of pubescence exists in this subspecies. Especially pubescent plants with small leaves occur in Chi- huahua and northern Durango. High montane plants tend to be shorter and more tufted, while the stems of low altitude plants are longer and more erect. Broader leaves occur in the latter populations, especially where they intergrade with C. hartwegii subsp. maccartii in northeastern Mexico and in western Texas, an extensive zone occupied by intermediates between subspp. hartwegii and pubescens. In one field study, D. E. Breedlove (personal communication) found anthesis in a population of this form to occur by 1 to 1% hours after sunset (San Luis Po- tosí, Breedlove 15473). However, no insects visited the flowers during the period of his observations. In the state of Chihuahua, Mexico I observed several bees ( Agapostemon, Apis) and small butterflies on freshly opened flowers in the midafternoon (Towner 247). Anthesis in Chihuahuan populations occurred from 4% to 2 hours before sunset. Greenhouse-cultivated plants from a wide range of Mexican localities showed great variation in opening times, extending from 4% hours before sunset to sunset. This variation in anthesis times may be a result of latitudinal or seasonal differences in photoperiod or a product of adapta- 1977] TOWNER—CALYLOPHUS 75 tion to locally differing insect faunas. Regional ecological differentiation may well have occurred within this subspecies since the southern forms from pine Forests seem to open much later than those inhabiting scrub and grasslands in the plains and hills of Chihuahua. Hawkmoths are undoubtedly regular evening visitors to all populations of subsp. hartwegii. Ultraviolet-absorbing areas on the petals can be nearly absent, present along the basal portion of the veins, or present as a small basal spot (Fig. 8). This suggests that some populations, i.e., those with spots absent, may be exclusively moth-pollinated while those with spots and early anthesis may be visited by bees active in the late afternoon. Chromosomal variation in this subspecies includes polyploidy, translocation heterozygosity, and extra chromosomes. Two of 11 populations, 1 from Aguas- calientes and 1 from Zacatecas, yielded tetraploid counts. Translocation heterozy- gosity was found in 7 of 17 diploid plants examined and in 6 of 9 populations from which meiotic determinations were obtained. The mean frequency of translocations per plant, 0.4, was the lowest for any taxon in the genus. Four plants of this subspecies had 1 to 4 extra diminutive chromosomes. Examination of hybrids indicated that one population of C. hartwegii subsp. hartwegii differed from the other taxa in sect. Salpingia by one or two translocations. Introgression of C. hartwegii subsp. hartwegii with other taxa appears to oc- cur widely, especially with subspp. pubescens and maccartii. Intermediates be- tween subspp. pubescens and hartwegii exist in southern West Texas, southeastern Arizona, and probably in northern Chihuahua and Coahuila. Regions of inter- mediate altitude in northeastern Mexico and along the upper Rio Grande River in southern Texas contain populations varying on a continuum between subspp. maccartii and hartwegii. As mentioned on p. 99, C. tubicula subsp. strigulosus may represent a stabilized derivative of introgression between C. tubicula subsp. tubicula and C. hartwegii subsp. hartwegii. Lastly, plants with narrow leaves, but lacking dense strigose pubescence, occur near Saltillo in southern Nuevo León and may represent introgressants with C. hartwegii subsp. filifolius, which occurs to the south of that area, or alternatively they may be independent nar- row-leaved derivatives of the species. No examples of sympatry without hybridization have been documented for C. hartwegii subsp. hartwegii and other taxa. The recent discovery of C. lavan- dulifolius in southern Nuevo León opens the possibility that it might come into contact with C. hartwegii subsp. hartwegii. Those two taxa proved somewhat intersterile in laboratory crosses, and might not be expected to hybridize exten- sively in the field. Ib. Calylophus hartwegii ( Benth.) Raven subsp. maccartii (Shinners) Towner & Raven, Madrono 20: 243. 1970 Calylophus hartwegii (Benth. ) Raven 2 IM Shinners, ens 1: 343. 1964. Oenothera greggii A. Gray var. pringlei Munz sensu Munz, A J. Bot. 16: 711. 1929, pro parte. O. pringlei (Munz) Munz sensu Munz, N. Amer. "FL ser. 2, 5: 138. 1965, pro parte. Stems several to many, sparingly branched above, nearly prostrate to ascend- g, 0.5-5 dm high; plants glandular-pubescent or minutely strigulose. Leaves 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 sparse to dense on stems, spreading to more or less ascending, narrowly lanceolate to lanceolate or oblanceolate, rarely linear, 6-35 mm long, 1-6 mm wide, nearly glabrous to sparsely strigulose or glandular-pubescent, the tip acute, the base acute-attenuate, the margin subentire to serrulate, usually undulate or undulate- crinkled; axillary leaves present, to 15 mm long. Inflorescence sparsely to densely strigulose. Floral tube 17-45 mm long, 5-12 mm wide at the throat. Se- pals 11-27 mm long, 2.5-7 mm wide, with free tips 1-6 mm long, occasionally with purple marginal stripes. Petals nearly orbicular to squarish, 10-30 mm long, frequently fading purple or pinkish, with a conspicuous, large basal ultraviolet- absorptive spot. Filaments 6-12 mm long; anthers 5-9 mm long. Style 25-60 mm long, glabrous above to minutely pubescent basally; stigma 2-4 mm broad; ovary 5-15 mm long, 1.5-2 mm wide. Capsule 10-22 mm long, 2-3 mm wide; seeds 1-2 mm long. Self-incompatible. Gametic chromosome numbers, n — TYPE: UNITED STATES. TEXAS: Starr Co., U.S. Highway 83, 6 mi NW of Rio Grande, in mesquite savannah, 24 March 1963, Rosa Ena Benavides 91 (SMU, holotype; TEX, isotype). Distribution (Fig. 15): Common semiarid grassy flats, in sandy to gravelly soil, often of limestone, frequently with Prosopis glandulosa, Opuntia, Acacia, Larrea divaricata, and Yucca, on the South Texas Plains and along the Rio Grande from Val Verde, Kinney, Uvalde, and Milam cos., Texas, south to southeastern Coahuila, central Nuevo León and northwestern Tamaulipas. From elevations of ca. 30 m (4 mi NW of Mathis, San Patricio Co., Texas) to ca. 1,500 m (Saltillo, Coahuila). Flowers March to September. Representative specimens 5 TEXAS: Dimmit Co.: E of Carrizo Springs, Jones 28153 (MO, POM). 8 mi S of Catarina, Ria id 16774 (DS, KANU), Duval Co.: 10 mi SW of Piece García 113 (OKLA, SMU, TEX). 16 mi NE of Freer, Malacara & Gutiérrez 30 (LL, U). San Diego, Tharp ur (TEX, US). 7 mi E of Freer, Rodríguez 104 (OKLA, SMU, TEX). Goliad Co.: Goliad, Williams 110 (PH, TEX). Jim Hogg Co.: 2 mi N of Santa Elena, Ríos & Cavazos 68 (LL). n bre Co.: 23 mi N of Alice, Painter vi al. 14436 (LL, TEX). 8 mi N of Alice, Bruni et al. 13 (LL). 15 mi NW of Alice, Castillo 20 (DS, SMU). Kinney Co.: 5 Treleau in s. (MO). Ca. 20 mi EE of Brackettville, Strother 299 (SMU, TEX). 26.0 mi SE of i Rio, Towner 34 (DS). La Salle Co.: Encinal, Vásquez 43 (DS). Live Oak 5 11.5 mi S of George West, Cory pr (POM). 8 mi S George West, Flyr 353 (DS, SMU). ur Co.: 30 m i SW of Eagle Pass, Bruni 8 (LL, OKLA, SMU, TEX). 5 mi N of Eagle Pass, Rowell 8894 (LL, OKL, OKLA). Eagle Pass, Schott in 1852 (F). San Patricio Co.: 4 mi NW of Mathis, Raven & Gregory 19386 (DS). Uvalde Co.: 5 mi of Uvalde, Munz 15558 (POM). Sabinal, 8165 29563 (POM). Val Verde Co.: Near Comstock city limits, Warnock & Turner 696 (SMU). N of Del Rio, Jones 28158 (MO, POM). Devil’s River, Earle & Earle 441 (MO, NY, US). Ca. 20 mi NNW of Del Rio, McVaugh 8259 (DS, F, SMU, TEX). Ca. 23.5 mi abf of Del Rio, Towner 32 (DS). 3.4 mi SE of Del Rio, Towner 33 (DS). Webb Co.: Minera, Reverchon 3558 (MO, US). 10 mi S of po Cisneros 15 (LL, OKLA). Laredo, T 6444 (LL, US). 11 mi S of Laredo, Robles 1 (SMU). 8 mi NW of Laredo, Ramírez 45 (DS, SMU). 23 mi PM of Laredo, McCart 225 (OKLA). 9.5 mi S of Laredo, Cory 28118 (POM). Zapata Co.: Zapata, Pérez 42 (DS). Near Zapata, Wood 42 (TEX ). 5 mi S of San Ignacio, ee 27 (SMU). 3 mi S of Zapata, Sdnchez 85 (OKLA, TEX). Zapata, Guajardo 32 (LL, SMU). 2 mi SE of Zapata, o E 92 (LL, OKLA, SMU). co. TAMAULIPAS: Along the river road, 20 mi E of the International Highway, Escalant c 55 (SMU, TEX, OKLA). 3 mi SW of * Loreto Ran 1 rutchfield & Johnston. 5568A (TEX). 50 mi SE of Nuevo Laredo, Garcia & Garcia 35 (D IU ). NUEVO LEON: 24 mi W of Monterrey, Waterfall & Wallis 13214 13215 (RSA, B Monterrey, 1977] TOWNER—CALYLOPHUS 77 Fisher 272 (MO, US). Rio Santa Catarina, Monterrey, Arséne 6306 (MO, US). 65 mi S of Nuevo Laredo, Pus & Frye 2369 (DS, MO, NY, RM, RSA, SMU, UC, WTU). 9 mi S of Nuevo Laredo, Frye & Frye 2390 (NY, RSA, UC, US, WTU). 12 mi N of Sabinas WI mu > Heard 14535 (F, MO, US). 17 mi NE of Sabinas Hidalgo, Rodríguez 70 (SM TEX 6 km W of Sabinas Hidalgo, Domínguez & McCart 8255 (SMU, TEX). 45 mi : of d de McCart et al. 8133 (OKLA, SMU, TEX). Sabinal (?), Jones 29563 (MO, UC). errey, Dodge T (US). Between Monterrey a nd Reynosa, along side road to San Juan, Langman 2870 (DS, PH). Monterrey, Edwards d» Eaton in 1846 (NY). 50 mi S of Laredo, less & Hall 637 (OKL). Ca. 54 mi : of “w U.S. border in Laredo, Towner 35 (DS). 39 mi N of center of Monterrey, Towner 36 (DS). COAHUILA: cm S of Parras, Sierra Negra, Stanford et al. 158 (ARIZ, MO). Near Diaz (now Piedras Negras ), r 55 8304 (DS, F, MO, PH, POM, RM, RSA, UC, US). Ciudad de Porfirio Díaz, Canby 109 (US). Guadalupe, Aguirre 703 (RSA). Porn. Aguirre & Reko 82 (NY). Ca. 48 mi N of Saltillo, Jackson 6722 (KANU). 25 km S of Piedras Negras, Rinehart 218 (OKL, OKLA, RSA). 13.4 mi S of central Saltillo, Towner 52, 53 (DS). — Closely related to Calylophus hartwegii subsp. hartwegii, this subspecies oc- curs at higher latitudes and lower elevations. It is relatively common in dis- turbed areas in the grasslands of southern Texas. It corresponds closely to var. maccartii as treated by Shinners except that narrower-leaved plants are included here. The broad leaves, which are frequently oblanceolate, early afternoon anthe- sis, and sparser, often glandular pubescence serve to distinguish this subspecies from subsp. hartwegii. Only leaf dimensions serve adequately in separating subspp. maccartii and filifolius. Considerable phenotypic variation occurs in subsp. maccartii in terms of pubescence, leaf shape, stature, and nature of the leaf margin. Leaf margins may be serrulate, undulate, or subentire. Pollination was studied near Saltillo, Coahuila, Mexico (Towner 52, 53) at a roadside population of C. hartwegii subsp. maccartii. Anthesis was not ob- served, but had been completed by 2% hours before sunset. Most visitors to the flowers were halictid bees, especially Evylaeus and Agapostemon, some of them possibly oligoleges. These may have played some part in pollination in the late afternoon and morning in spite of their small size. No large native bees, except for a single Bombus, and no hawkmoths were observed visiting flowers, but their involvement cannot be discounted on the limited evidence available. reenhouse studies showed anthesis times occurring 3-5 hours before sunset and flowers with large central ultraviolet-absorbing areas. Self-incompatibility was found in the 3 plants available for testing. This suggests that C. hartwegii subsp. maccartii has perhaps secondarily shifted from hawkmoth pollination, as indicated by its TAE to bee pollination, as might be inferred from its ultraviolet pattern and behavi Two of 10 populations howe tetraploidy, in addition to 1 population inter- mediate between Calylophus hartwegii subspp. filifolius and maccartii. One plant from each of 2 populations had a single extra diminutive chromosome. Half of 16 plants, representing 5 of 8 diploid populations, were heterozygous for trans- locations. The mean frequency of translocation heterozygosities was 0.6 per plant, and only one plant had as many as 2. Experimental hybrids between C. hartwegii subsp. maccartii and other members of sect. Salpingia were heterozy- gous for 1 or 2 reciprocal translocations. Introgression occurs, as mentioned above, with C. hartwegii subsp. hartwegii in southern Texas and northeastern Mexico. It also appears to have taken place 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 with subsp. pubescens along the upper Rio Grande in southern Texas, although there is difficulty in recognizing the sources of variation in this region. Two further collections [36 mi W of Monterrey, Coahuila, Mexico, Towner 39 (DS). 5 mi W of Marathon, Brewster Co., Texas, Warnock 60004 (TEX)] appear to be intermediate between subspp. maccartii and filifolius, although this may not necessarily have resulted from introgression. The only other taxon of Calylophus occurring near C. hartwegii subsp. maccartii is C. berlandieri subsp. berlandieri. The two are essentially intersterile, but in the South Texas Plains have often been mistaken for one another because there they tend to resemble each other in leaf shape and character of the margin. lc. Calylophus hartwegii (Benth.) Raven subsp. filifolius ( Eastw.) Towner & Raven, Madroño 20: 243. 1970 Oenothera tubicula A. Gray var. filifolia Eastw., Proc. Calif. ne s ser. 3, 1: 72. 1897. Galpinsia 1 (Eastw.) Heller, Cat. N. Amer. Pl., ed. 2. 8. 1000. Oenothera ha rt- wegii Benth. var. filifolia (Eastw.) Munz, Amer. J. ` Bot. 16: 707. 1929. Calylophus hartwegii (Benth. ) Raven var. filifolius (Eastw.) Shinners, Sida 1: 345, 1965. Oenothera hartwegii var. fendleri (A. Gray) A. Gray subvar. filifolia (Eastw.) H. Lév. Monogr. 8 Onoth. 335. ; Oenothera hartwegii Benth. var. typica sensu Munz, Amer. J. Bot. 16: 706. 1929, pro parte. Calylophus hartwegii ( Benth.) Raven var. hartwegii sensu Shinners, Sida 1: 342. 1964, pro parte. Oenothera hartwegii var. hartwegii sensu Munz, N. Amer. Fl, ser. 2, 5: 139. 1965, pro parte. Stems several to many, moderately to densely branched above, decumbent and spreading to somewhat ascending, 0.5-4 dm high; plant minutely glandular- pubescent throughout, more densely so on the ovary and inflorescence, infrequently sparsely strigulose on the ovary and leaves. Leaves moderately well- spaced to dense on the stem, spreading to ascending, filiform to narrowly lanceo- late, 3-45 mm long, 0.4-3(-4) mm wide, the tip acute, the base acute-attenuate, the margin entire to remotely serrulate, occasionally undulate; axillary leaves present, to 10( +) mm long. Floral tube 16-50 mm long, 4-14 mm wide at the throat, occasionally fading pinkish. Sepals 7-17 mm long, 3-7 mm wide, with free tips 0.5-4 mm long, frequently with purple spotting and occasionally with a purple marginal stripe. Petals suborbicular to somewhat rhomboidal, 12-23 mm long, occasionally fading pinkish, with a basal ultraviolet-absorptive spot of moderate to large size. Filaments 6-13 mm long; anthers 6-11 mm long. Style 26-60 mm long, glabrous above, glabrous or minutely pubescent basally; stigma 1.5-4 mm broad; ovary 4-13 mm long, 1-2 mm wide. Capsule 7-22 mm long, 2-3 mm wide; seeds 1.2-2 mm long. Self-incompatible. Gametic chromosome number, n — TYPE: UNITED STATES. NEW MEXICO: White Sands, probably in Otero Co., August 1896, T. D. A. Cockerell ( CAS). Distribution (Fig. 16): Highly local, but often abundant, almost always on semiarid gypsum flats, dunes, or outcrops, frequently with Larrea divaricata, Yucca, or Juniperus, from Otero and Torrance cos., New Mexico south and east through the Trans-Pecos and southern Panhandle regions of Texas, thence north- east to Cottle Co., Texas and southward from widely scattered localities in cen- 1977] TOWNER—CALYLOPHUS 79 tral Chihuahua and Coahuila. Occurring from elevations of ca. 600 m (7 mi N of Spur, Dickens Co., Texas) to ca. 1,850 m (7.2 mi SE of Willard, Torrance Co., New Mexico). Flowers May to October. Representative specimens examined: JNITED STATES. NEW MEXICO: Chaves Co.: 20 mi S of Roswell, Earle & Earle 293 (MO, NY, POM, RM, UC, US). 2 mi E of Bottomless Lakes State Pack Headquarters, Hess 73 (WTU ). 56. 7 mi SE of Vaughn, Towner 125 ( DS). 18.7 mi N of Roswell on U.S. 285, Towner 128 (DS). De Baca Co.: 22 mi S of Fort Sumner, Brooks d» 1 25763 (DS). Dona Ana Co.: Jornada Game Reserve, Wooton, no date (U S). Eddy Co.: 3 mi NW of Texas state line on U.S. 62/180, Raven & p 191 56 (DS). 6 mi SW of White' 8 City, Munz & Gregory 23357 (POM, UC, WTU). 11.4 mi SW of White’s City, Towner 22 (DS). Lea Co.: 55-60 mi E of Roswell, Palmer 62 (F). 1 Co.: White Mts. , 5,400 ft, Wooton 181 (ARIZ, DS, GH, MO, NY, POM, RM, UNM, UC, US). 22 mi N of Cartizose: Brooks & Stephens 25957 (DS). Otero Co.: Round Mt., along Tularosa Creek in Sacramento Mts., Wooton in 1899 ( ARIZ, DS, NMC, NY, POM, RM, US). White Sands National Monument, Munz € Gregory 23335 (POM). White Sands National Monument, 2 mi W of headquarters, Toude 1l (progeny, DS). Torrance Co.: Near Willard, Wooton 2730 (COLO, DS, RM, US). 7.2 mi SE of Willard, Towner 122 (DS). TEXAS: Culberson Co.: 2 mi SE of U.S. 62/180 at New Mexico line, MoVeugh 8164 (DS, GH, LL, SMU, TEX). 30 mi N of Van Horn, Waterfall 4122 (GH). | Co.: 7 mi N of ns Moss 19 (OKLA). Ector Co.: 1 mi E of jct. of Texas 185 and U.S. 385, ae 424 (DAO, DS, RSA, UC). Gaines Co.: 15 mi E ad Seminole, Lundell & Lundell 16955 (LL). Howard Co.: Big Springs, Tracy 8306 (F. GH, MO, NE B, NY, TEX, US). Hudspeth Co.: Gypsum quarry E of Finley, Waterfall 5023 (G H, MO). Kent Co.: 2 mi bd of Clairemont on U.S. 380, Correll & Johnston 22107 (LL). Towing Co.: ibas Salt eek near zwar in N of Orla, Correll & Correll 26016 (Mixed with C. hartwegii subsp. 5 LL). N n Co.: E of Stanton, Lundell & Lundell 16916 (LI) Midland Co.: 4 mi E of Midland 1 12000 (POM, TEX). Nolan Co.: Sweetwater, Reverchon 1285 (F, MO). Ward Co. 9.5 mi S of Monahans, Gregory 174 (RSA, UC, W TU). Winkler Co.: 1 mi N of southern county 15 5 on highway 18, Irving 69 (SMU, TEX). MEXICO. COAHUILA: Morillo, Saltillo, Lyonnet 3497 (US). Saltillo, Fisher in 1926 (DS). 6 mi N of La Ventura, Johnston 7644 d Shreve 8726 oe UC, US). NUEVO LEÓN on COAHUILA: Vanegas-Saltillo road, alkali plain, Lundell 5721 (ARIZ, F, POM 1). CHIHUAHUA: 13 mi S of Gallegos, Breedlove 15734 (DS). zacatecas: Intersection of highways 49 and 45, Cruden 1238 (TEX). No locality: Mexico, Gregg 33 (MO). — Calylophus hartwegii subsp. filifolius, which is largely endemic to gypsum soils, may include some convergent populations of independent origin. The Texas and Coahuila populations are widely separated, with no collection having yet been obtained in the intervening region, a span of over 650 I have retained here certain populations with broader leaf SPEM than were included by Shinners or Munz. Plants from the type locality do not all have filiform leaves, although collections from the White Sands area do include the narrowest-leaved plants in the species. Inclusion of the broader-leaved popula- tions here simplifies the variation pattern in C. hartwegii subsp. hartw egii and renders subsp. filifolius a major geographical race which is nonetheless phenet- ically discrete. In West Texas and southeastern New Mexico this taxon is abun- dant on plains at about 600-1,200 m elevation. Some variability is shown by this subspecies in terms of leaf width, petal shape, the presence and distribution of anthocyanins, and length of the free sepal tips. Pubescence is relativ 'ely uni- form, with nearly all plants being glandular-pubescent, and only a few being even minutely strigulose. The pollination studies of Gregory (1964; as Oenothera hartwegii) in Ector and Ward cos., Texas indicated that flowers of C. hartwegii subsp. filifolius were 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 { / É 7| Ü { | o " ) ` enn be x ] | — | | = | | s | | | | | ! | ° œ — J — ° ‘oe E = 2. x ^ | o e 5 0 ° N 8 š MP a 8 ; s |... m ` o Je ° + s S 4 eu | ik 1 * 2 ° f 9 e? e 8 E "s oe e e ig Ny C z e | g3? 22 I „ „% ° — | ° ° °% 5. P a ° | Ld 4 ae à 20 cee oe | ° thee IE aat i T °° ° ee e.a ° 2 oe ew ° * ° — ° ° ° ° e \ oo ee , ee \ E e ° * ° | y [E ee, A E ^ e e oa VO B 4 D Fic 16. Distributions of Calylophus hartwegii subsp. pubescens (dots), C. hartwegii subsp. una (open circles), and C. hartwegii subsp. filifolius (triangles). 1977] TOWNER—CALYLOPHUS 81 open well before sunset and were visited by hawkmoths in the evening and sometimes by bees in the afternoon. Greenhouse studies showed anthesis to oc- cur 3-6 hours before sunset. Ultraviolet-absorbing spots at the base of each petal were fairly large, presenting conspicuous regions of high contrast which would be visible to diurnal insects. Tests for self-incompatibility on 6 plants all prove positive. Cytogenetically, C. hartwegii subsp. filifolius exhibits a great deal of varia- tion, with plants averaging 1.7 translocation heterozygosities. Ten of 12 plants from 8 populations were heterozygous. The maximum association of chromo- somes observed consisted of a ring of 12 and a bivalent, present in a plant grown from seed collected in Winkler Co., Texas (Irving 69). As many as ll extra diminutive chromosomes were observed in plants from that population, with some being possessed by each of the 4 plants examined. No other population demonstrated extra chromosomes, and no tetraploid or higher counts were ob- tained from this subspecies. One possible intermediate between C. hartwegii subspp. maccartii and filifolius, as mentioned before, was tetraploid (Towner 39). Crosses of C. hartwegii subsp. filifolius with other forms in sect. Salpingia demonstrated complete homology with C. tubicula, C. hartwegii subsp. fendleri, and C. hartwegii subsp. pubescens, and one or two translocation differences from the other taxa. Calylophus hartwegii subsp. filifolius intergrades somewhat with C. hartwegii subsp. fendleri in the southern Texas Panhandle and in New Mexico, but hybridi- zation is limited by the altitudinal separation of these subspecies. Similarly, there is limited hybridization with C. hartwegii subsp. pubescens in the same regions. Possible hybridization with C. hartwegii subsp. maccartii and C. hartwegii subsp. hartwegii was treated under those taxa. Sympatry without hybridization occurs in New Mexico where C. berlandieri and C. serrulatus occasionally come into contact with this subspecies. ld. Calylophus hartwegii ( Benth.) Raven subsp. fendleri (A. Gray) Towner & Raven, Madroño 20: 243. Oenothera fendleri A. Gray, Mem. m Acad. Arts. , 4: 45. 1849. O. hartwegii ieu leri (A. Gray) A. Gray, Pl. Wright. 2: 58. 1853. Galpinsia hartwegi ( Benth.) Britton (var.) fendleri (A. Gray) Small, Bul. Torrey Bot. Club 23: 186. 1896. G. a . Gray) Heller. Cat. N. 1900. Oonothera hartwegii Benth. var. ee sensu p Amer. J. Bot. 16: 706. 1929, pro parte. Calylophus hartwegii ( Benth.) Raven var. hartwegii sensu Shinners, Sida 1: 342. 1964, pro parte. en iothera hartwegii var. hartwegii sensu Munz, N. Amer. Fl., ser. 2, 5: 139. 1965, pro par Stems one to several, sparingly to moderately branched above, ascending to more or less erect, 1.5-4 dm high; plant glabrous throughout, infrequently mi- nutely and sparingly glandular-pubescent. Leaves sparse to dense on stems, more or less ascending, linear to oblanceolate or lanceolate, 10-50 mm long, 1.5-10 mm wide, the tip acute, the base acute-attenuate to obtuse, infrequently nearly clasping, the margin entire to subentire, infrequently undulate; axillary leaves usually absent, to 10 mm long when present. Floral tube 30-50 mm long, 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 7-15 mm wide at the throat, sometimes with purple lines, frequently fading pur- plish or orange. Sepals 9-28 mm long, 4-10 mm wide, with free tips 0.5-3 mm long, occasionally with purple margins or spotting. Petals obovate to somewhat rhomboidal or squarish, 10-30 mm long, usually fading purplish or reddish, with a basal ultraviolet-absorptive spot small or absent. Filaments 5-12 mm long; anthers 5-13 mm long. Style 40-75 mm long, glabrous above, minutely pubescent basally; stigma 2-6 mm broad; ovary 7-20 mm long, 1-2 mm wide. Capsule 10-40 mm long, 2-3 mm wide; seeds 1-1.5 mm long. Self-incompatible. Gametic chromosome number, n — 7 TYPE: UNITED STATES. NEW MEXICO: without specific locality, "prob- ably Santa Fe,” 1847, A. Fendler 230 (GH, lectotype; GH, MO, NY, P, PH, US, isolectotypes); cf. Munz, Amer. J. Bot. 16: 708. 1929. The sheets assigned this number are probably a mixture of collections from the originally published lo- calities, i.e., Santa Fe, on the Río del Norte (Rio Grande), and from Rock Creek eastward to the Cimarron River. Distribution ( Fig. 16); Uncommon on clay or gravelly soils, occasionally cal- careous, from high plains with Prosopis glandulosa and Juniperus, to montane forests with Juniperus, Pinus edulis, and occasionally Pinus ponderosa, from Bar- ber and Morton cos., Kansas, south through western Oklahoma and widely scat- tered sites in the Texas Panhandle to eastern Chihuahua, central Trans-Pecos Texas, central and western New Mexico, and east-central Arizona. Elevational distribution from ca. 370 m (Agawam, Grady Co., Oklahoma) to 2,200 m (17 mi N of Alpine, Apache Co., Arizona). Flowers April to October. Representative specimens examined: UNITED STATES. KANSAS: Barber Co.: NW corner of county, Baker in 1904 (NY). Gypsum hills, Hitchcock 689 ( GH, NMC, NY, RM, US). 6 mi W of Medicine Lodge, 2e ns 11150 (KANU, OKLA). 7 mi W of Medicine Lodge, McGregor 14243 (KANU, SMU, 7 mi SW of Medicine Lodge, McGregor 14472 (SMU, US). Sandy soil S of Coats, CER in 1936 ( ARIZ, CAN, F, MO, OKL, OKLA, RM). 7 mi S of Sun City, McGregor 14019 ( KANU ). Morton Co.: 4 mi W of Rolla, McGregor 12858 (KANU, SMU, US). OKLAHOMA: Blaine Co.: mi NE of Watonga, Stephens & Brooks 20819 (KANU). Roman Nose State Park, Goodman & Waterfall 4185 (GH, OKL, OKLA). Grady Co.: 8 mi SW of Chickasha, Pearce 767 (SMU) Greer Co.: 7.7 mi S of Mangum, Towner 79 (DS). 2.6 mi S of Mangum, Towner 85 (DS). 0.5 mi S and 4.3 mi W of Brinkman, Towner 86 (DS). Harmon Co.: 10 mi S of McQueen, Stephens & Brooks 20758 (DS, KANU). Harper Co.: Near Buffalo, Stevens 535 (DS, GH, O, NY, OKL, OKL A, US). 17 mi E and 7 S of Buffalo, Stephens & d 21685 (DS, q Kiowa Co.: Snyder, Stevens 1198 (OKLA). Roger Mills Co.: 18 mi N of Cheyenne, Waterfall 11897 (OKLA, SMU, TEX, US). Ca. 8 mi E of Strong City, Toner 156 (DS Co.: 37 mi W « X Alva, Stratton 6384 (KANU, OKL). Woodward C 24 mi N of Mooreland, Brooks d» Ste phens 21658 (DS, KANU). texas: Hemphill Co: Pr rairies N of Canadian, Eggert in 1901 (MO). Jeff Davis Co.: 14.8 mi N of Marfa city limits, Parnell 68-T-30 ( DS). 8 mi S of Fort Davis, Munz & Gregory 23384 1 UC). 3.8 mi W of Fort Davis, Gregory 134 (NY, RSA, WTU). Mesa S of Fort Davis, Andrews 63 (COLO, GH). Limpia Canyon, Davis Mts., Bray in ey (TEX). Presidio Co.: 8 mi E of Marfa, Warnock 7916 (LL, SMU, TEX). 8 mi NW of Marfa, Jackson in 1964 (progeny only, DS). 13.0 mi W of Marfa, Towner 25 (DS). 11.9 mi NW of Marfa, Towner 26 (DS). 10.4 mi NW of Marfa, Towner 27 (DS). Randall Co.: Bottom of canyon, Palo Duro Canyon State Park, Lundell & Lundell 11442 (LL, SMU). Wilbarger Co.: 1.5 mi S of Harrold, Whitehouse 9764 (ARIZ, SMU, UC, US). NEw Mexico: Catron Co.: Mangas Creek, Rusby in 1880 (US). Mangas Canyon, Greene in 1880 (F, MO, NY, PH). Grant Co.: Vicinity of Gila, yond 11630 (DAO, NY, WTU). Lincoln Co. 2j: Callinas Mts., Wooton 2741 (US). Oter 8 mi E of Mescalero, Parker 2556a ( ARIZ, COLO). Above Masedlaim. White Mts., Wooton in ~ 1977] TOWNER—CALYLOPHUS 83 1895 (US). Rio Arriba Co.: Arroyo de Agua, Gregory 588 (UC). Sandoval Co.: San Isidro, Benedict 2311 (US). San Miguel Co.: Near Pecos, Standley 4952 (GH, MO, NMC, NY). Santa Fe Co.: 19 mi M of Santa Fe, along the Rio Grande R., Heller d» Heller 3622 (MO, Y, US). Socorro Co.: 0.4 mi S of Magdalena, Towner 119 (D S). Torrance Co.: 3.1 mi NW of Cedarvale, Toone 123 (DS). 2.6 mi W of Willard, Towner 121. Valencia Co.: 8 mi E of Ramah, Wooton in 1906 (NMC, NY, US). arizona: Apache Co.: 17 mi N of Alpine, Breedlove 14298 (DS). 7.9 mi N of Alpine, Towner 110 (DS). 4.5 mi N of Nutrioso, Towner 111 (DS). 80 mi N of Nutrioso, Towner 112 (DS). Coconino Co.: Walnut Canyon, MacDougal in 1898 (ARIZ, F, GH, NY, PH, RM, UC, US). Flagstaff Cemetery, Demaree 42847 ( ARIZ, DS, OKLA, RSA, SMU). Navajo Co.: Near Heber, Parker et al. 6832 (ARIZ, F). > mi N of Whiteriver, Goodman & Hitchcock 1298 (DS, F, MO, NY, PH, UC). MEXICO. CHIHUAHUA: ll mi E of Highway 16 on road to new lake on Rio Conchos, Powell et al. 2082 (TEX). A race with distinctive characters of distribution, morphology, and floral behavior, Calylophus hartwegii subsp. fendleri definitely merits recognition, contrary to the opinion of Shinners (1964). Late anthesis, glabrous vegetative parts, and distribution at relatively high altitudes or latitudes are strongly cor- related in this form. In the northern part of its range, it occurs at intermediate or low altitudes, but in Arizona and New Mexico it ranges up into the coniferous forests. Leaf dimensions not being of critical importance for delimiting this subspecies has allowed the inclusion of elements from Munz's Oenothera hart- wegii var. hartwegii. Thus considerable variation in leaf dimension is retained in C. hartwegii subsp. fendleri, which attains a large and broad leaf size for the species, especially in collections from the Great Plains. Relatively narrow-leaved forms from the mountains of Trans-Pecos Texas belong here, although they have been traditionally placed with Oenothera hartwegii var. hartwegii. The type series of Evylaeus galpinsiae ( Cockerell) was collected from Caly- lophus hartwegii subsp. fendleri near Pecos, New Mexico where the bees were active at 7:30 in the evening (Cockerell, 1903). Pollination studies of Gregory (1964) in Jeff Davis Co., Texas (as Oenothera hartwegii) suggested that anthesis occurs near sunset and that pollination is largely accomplished by hawkmoths. My field observations in Grant Co., New Mexico (Towner 244) differed some- what from Gregory's in that numerous bees of the genera Sphecodogastra and Evylaeus were active gathering pollen from this subspecies for about one hour starting at sunset. As observed by Gregory, hawkmoths visited the flowers heavily in the early evening. Infrequent visits by bees were seen in the morning. Field observations at this site and elsewhere in New Mexico indicated that anthesis occurs at about sunset, and flowers on cultivated plants opened within an hour before or after sunset. Photography under ultraviolet light showed plants to have either no spots of absorption on the petals or only very small ones ( Fig. 10). Two plants were checked and found to be self-incompatible. No tetraploid individuals have been found in this subspecies. Sixteen of 23 plants from 12 of 15 populations were heterozygous for translocations, with a mean number of 0.9 heterozygosities per plant. The maximum number of heterozygosities exhibited was 2, seen in 5 plants. Hybrids with other taxa in sect. Salpingia behaved identically to those involving C. hartwegii subsp. fili- folius in terms of chromosome pairing. Four to 5 extra diminutive chromosomes were observed in 3 plants, each from a different population. Instances of hybridization with C. hartwegii subsp. pubescens, particularly in 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Oklahoma, seem relatively common, and the two subspecies are not separated by large ecological differences. One population in Harmon Co., Oklahoma consisted of both subspecies and intermediates. Cases of intergradation with other taxa have been discussed in previous sections. Calylophus hartwegii subsp. fendleri occurs frequently with C. serrulatus in Oklahoma, with no indication of hybridi- zation. A population in Torrance Co., New Mexico was found growing with C. lavandulifolius, and again no intermediates or putative hybrids were observed. le. Calylophus hartwegii ( Benth.) Raven subsp. pubescens (A. Gray) Towner & Raven, Madroño 20: 243. 0. gii em ae A. ~ var. pubescens A. Gray, Pl. Wright. 1: 72. 1852. ee hartwegii (Benth.) Raven var. pubescens (A. Gray) Shinners, Sida 1: 344. 196 Sn. greggii A. Gray, Mem. Amer. Acad. Arts, n.s., 4: 46. 1849. Galpinsia sage ( A. Gray) Small, Bull. oce Bot. Club 23: 186. 1896. Oenothera hartwegii Benth. var. dap) (A. Gray) A. Gray iier filifolia H. Lév. f. thymifolia H. Lév., Monogr. Onoth. 5. 1908, based on MO isoty O. greggii var. 1 Munz, Amer. J. B n 16: 709. 1929. ryPE: Mexico, Durango, hill SE of Pelayo, ca. 60 mi NW of Torreón, 8 May 1847, Josiah Gregg 591 (GH, holotype; MO, NY, isotypes). m lampasana Buckley, Proc. Acad. Nat. Sci. Philadelphia 1861: 454. 1961. 5 mpasana (Buckley) Wooton & Standley, Contr. U.S. Natl. Herb. 16: 152. 1913. Oenothera reget A. Gray var. lampasana (Buckley ) Munz, Amer. J. Bot. 16: 710. j. TYPE: ed States, ex Lampasas Co., prairies, 1860-1861, S. B. Buckley ( PH). Galpinsia interior Small, Fl. S.E. U.S. 845, 1335. 1903. rype: United States, d igi Co., Fort Niobrara, 25 io 1888, T. E. Wilcox (NY). This locality is more thar mi » of te known range of this subspecies and probably resulted from dispersal by 1 r human intent, or the label may have been switche Gala camporum Wooton & Standley, Contr. U.S. Na d. Herb. 16: 152. 1913. Oenothera , (Wooton & Standley) Tidestrom in Tidestrom & Kittell, Fl. Ariz. & N. Mex. 278. "1941. TYPE: United States, New Mexico, Lea Co., Knowles, 29 July 1860, E. O. Wooton (US-564592, holotype; NMC, POM, US, isotypes s). Stems several, moderately branched above, decumbent to more or less erect, 1-5 dm high; plant usually covered throughout with long spreading trichomes, most densely on the ovary, inflorescence, and upper stem, occasionally also with short glandular or nonglandular trichomes. Leaves somewhat sparse to dense on the stem, most commonly spreading to reflexed downward, sometimes more or less ascending, very narrowly elliptic or narrowly lanceolate to ovate, 5-40 mm long, 1.5-12 mm wide, the tip acute, the base acute to truncate or subcordate and clasping, the margin entire to sparsely serrulate, occasionally undulate- crinkled; axillary leaves often absent or much reduced, occasionally to 15 mm long. Floral tube 20-50 mm long, 4-20 mm wide at the throat, only rarely with purple stripes, occasionally fading purplish. Petals obovate to somewhat rhom- boidal or squarish, 12-35 mm long, frequently fading pinkish or purplish, with a basal ultraviolet-absorptive spot of small or moderate size. Filaments 5-12 mm long; anthers 4-13 mm long. Style 25-70 mm long, glabrous above, minutely pubescent basally; stigma 1.5-5 mm broad; ovary 5-30 mm long, 1-3 mm wide. Capsule 6-35 mm long, 2-3 mm wide; seeds 1-1.7 mm long. Self-incompatible. Gametic chromosome numbers, n = 7, 14. TYPE: UNITED STATES. TEXAS: dry hills beyond the Pecos River, probably from Pecos Co., August 1849, Charles Wright 199 (GH, holotype; GH, NY, PH, 1977] TOW NER—CALYLOPHUS 85 US, isotypes). The locality was calculated from the dates and account of Wright's trip given by McKelvey (1955: 1067-1068 Distribution (Fig. 16): Common and colonial in modesta drv open places, plains, and hills, in sandy to gravelly soil, often of limestone or gypsum, fre- quently with Prosopis glandulosa and Juniperus, from Baca Co. and eastern Las Animas Co., Colorado and Morton and Meade cos., Kansas, to western Okla- homa and the Texas Panhandle, throughout central and Trans-Pecos Texas, thence west through eastern and southern New Mexico to central and southeastern Ari- zona; also south very locally in central Coahuila and northeastern Durango. Ele- vational distribution from 200 m (10.5 mi E of Weatherford, Parker Co., Texas) to 2,100 m (2.4 mi NW of Corona, Torrance Co., New Mexico). Flowers March to October. Representative specimens examined: UNITED STATES. COLORADO: Baca Co.: 20 mi S of Pritchett, Harrington 3325 (RSA). 27 mi S of Pritchett, Weber 4608 (COLO, UC, WTU). Las Animas Co.: 7 mi S and 16 E of Kim, Weber 4387 (COLO). 4 mi W of Andrix, Rogers 4952 (COLO, US). KANSAS: Clark Co.: 10 mi S of Ashland, Rydberg & Imler 744 (NY). Meade Co.: 8 mi S and 7 E of Meade, Horr in 1957 P SE corner of county, above Wolf Canyon, Horr 3612 (KANU). Morton Co.: Stony hills, Hitchcock 166 ( GH, MO, NMC, NY, POM, RM, US). Point of Rocks, Hitchcock 634 (GH). On Cimarron R., N of Elkhart, Point of Rock, Rydberg & Imler 9 2, 943 (MO, NEB, NY). Point of Rocks, 7 mi N and 4 W of Elkhart, Stephens 12955 (KANU). No county: SW Kansas, Plank in 1886 (GH). OKLAHOMA: Beckham Co.: 8 mi N of Sayre, Wiedman in 1959 (OKL, OKLA). 6 mi S of Elk City, Eskew 1503 (GH, KANU, OKL, OKLA). 1.7 mi W and 2.4 N of Elk City, Stratton 6835 (KANU, OKLA). Cimarron Co.: 16 mi SE of Kenton, Waterfall 7433 (OKL, TEX). 2 mi N of Kenton, Hopkins & van Valkenburgh 5754 (NY, RM, SMU). Custer Co.: mi and 0.3 S of Weatherford, Waterfall 442 (OKLA, POM). Canyon rims, Clinton, Demaree 12466 (ARIZ, GH, MO, NY, OKL, PH, POM, SMU, US). 10 mi W of Clinton, Munz & Gregory 23508 (RSA). Ellis Co.: Near Shattuck: Clifton 3174 (GH, OKLA). Greer Co.: 2 mi S of Mangum, Robbins 3038 (NY, OKL). 3 mi S of Mangum, Stephens 20812 (DS). 4.5 mi S of Mangum, Towner 82 (DS). Harmon Co.: Near Hollis, Stevens 1162 (DS, GH, MO, NY, OKL, OKLA, US). 13.5 mi W of Mangum, Waterfall 7174 (OKL, OKLA). Jackson Co.: 3 mi N and 1 W of Eldorado, uoc 9008 (OKL, OKLA). Kiowa Co. 3 mi W of Gotebo, Goodman 6274 (OKL, RSA, UC). Roger Mills Co.: Red lands, E 417, 418 (OKL). 2.5 mi S of Cheyenne, Wiedeman 183 (OKL, OKLA). Texas Co.: Goodwell, Butler 85 (OKLA). 5.5 mi E of Hardesty, Stephens & Brooks 21775 (DS). 7 mi NE of Texhoma, Waterfall 9123 (GH, OKL, OKLA). TEXAS: Brewster Co.: Glass Mts., Tharp 3629 (US). 4 mi S of Alpine, Munz & Geon 23395 (RSA, UC). Hot Springs area, Sperry 1732 (GH). 3.3 mi W of Alpine, Towner 28 (DS). Brown Co.: 8.2 mi S of oc Towner 63 (DS). . Ca. 17 mi SE of Abilene, Mode a 64-53 (DS). Co.: 2 mi SE of Bronte, Raven & Gregory 19279 (DS). Coleman Co.: 14.3 mi N of roles man, . ; O). — A ~ =e > zy e 2 Q o i Eo Concho Co.: 2.5 mi W of Eden, Munz & Gregory 23425 (RSA, UC, WTU). 2.8 mi N of Eden, Racen & Gregory 19277 (DS). Culberson Co.: 27 mi SW of White’s City, New Mexico Munz & Gregory 23364 (RSA, UC). 10 and 12 mi N of Van Horn, Waterfall 4095 ( ARIZ, GH, MO, NY). Victoria Canyon, Sierra Diablo, Correll & Rollins 23783 (LL). 25.6 mi SW of 189 City, New Mexico, Towner 23 (DS). Glasscock E 3 mi E of Garden City, Munz & Gregory 23420 (RSA, UC, WTU). Hardeman Co.: i N of Quanah, Stephens 20721 (DS). Irion Co.: 30 mi N of Barnhart, Raven & Gregory des (DS). Lampasas Co.: Lampasas, Reverchon 1302 (DS, F, MO, NY, PH, UC, US). Maverick Co.: Eagle Pass, Havard s.n. (US). Mills Co.: 0.9 mi N of center of Goldthwaite, Towner 70 (DS). Pecos Co.: 25 mi NW of Sanderson, Munz & Gregory 23405 (RSA, UC, WTU). Potter Co.: 3.1 mi N of U.S. 66 on Farm Road 1719, Towner 93 (DS). 11.3 mi N zd 3.9 W of central Amarillo, aa Ae (DS). Presidio Co.: Ca. 35 mi S of Marfa, Bunton Flats, Warnock 46621 (RSA ). Ca. 3 mi SW of Marfa, Hine kley 706 (LL). 12 mi N of Shafter, Scuddy 396 (OKLA). Marfa, 1 17341 (US). Real Co.: Leakey, Palmer 10149 (DS, PH, POM). Roberts Co.: 24 mi 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 S of Perryton, drip 2988 (KSC). Taylor Co.: Camp Barkeley, Tolstead 7065 (MO SMU, UC). Terrell Co.: 6.7 mi E of Sanderson, Raven & dud 19202 (DS). 63 mi E of Sanderson, is 275 (DAO, RSA, UC). 9.6 mi W of Dryden, Parks et al. 305 925 SMU). Uvalde Co.: Sabinal, Palmer 11514a (MO). By 9 R., McKelvey 1879 (G POM). Uvalde, Dobie m 1930 (TEX). Val Verde Co.: Pumpville turnoff, Warnock ie (LL, SMU). Ca. 5 mi W of Langtry, Warnock & Cameron 9937 (LL, SMU). Ward Co.: Near Monahans, Wheeler in 1938 (LL). Wheeler Co.: Ca. 0.5 mi W of Shamrock, m 88 (DS). Counties unknown: Near Mt. Carmel, Rio G nds Parry 369 (NY, PH). On the Rio Grande, Wright in 7 isa Between Pa and d Kagan in 1 5 (TEX, biochemical voucher ). MEXICO: Chaves Co.: Ca. 5 mi N of Roswell, Towner 126 (DS). 13.5 mi of Hope, oau 13 (DS). Ca. 8 mi E of Elk. Towner 12 (DS). De Baca Co N side of Fort Sumner, Shinners 20922 (SMU). Dona Ana Co.: Organ Mts., Waman in 1900 (NMC, POM, RM, US). Eddy Co.: Near Three Forks of Rocky Arroyo, Guadalupe Mts. Wilken 1734 (PH, US). 1.5 mi ENE of headquarters, Carlsbad Caverns National Park, Dole UC). Junction a Delaware Creek and Pecos R., Pope in 1835 (GH). Memorial Hospital, N end of Car Fs 155 Munz d» ‘Ga 23355 (RSA, UC). Lea Co.: 1-11 mi N of Hobbs, dein 2569 (ARIZ). 60 mi E of Roswell, Palmer 66 (F). Lincoln Co.: Ca. 15 mi W of oswell, Dunn 905 (RSA). 10 mi E of 5 Hitchcock et al. 4201 (DS, UC, WTU). 18 (Hondo?) Hill, Wooton in 1904, 1906 (NMC). Otero Co.: 9 mi NE of Alamogordo, Munz & Gregory 23337 (RSA, UC, WTU). Quay Co.: Ca. 9 mi W of Tucumcari, Towner 94 DS). 8 mi SW of Tucumcari, Shinners 21062 (SMU). 8 mi S of San Juan, Stephens & Brooks 25573 (DS). Roosevelt Co.: Portales Springs, Martin 784 J). Near Causey, Wooton in 1909 (NMC). Sierra Co.: Berendo Creek, Metcalfe 1574 (F, GH, MO, NMC, Y, POM, , US). Torrance Co.: 2.4 mi NW of Corona. Towner 124 (DS). Union Co.: Ca. 4 mi N of Moses, York & Rodgers 147, 149 (SMU, TEX). arroxa: Cochise Co.: 15 mi E of Bernardino, Benson 10284 (ARIZ, POM, UC). 6 mi NW of Chiricahua, Gould & Pultz 3155 (ARIZ, GH, UC). 6 mi W of entrance to Chiricahua National Monument, Gregory 408, 411 (DAO, DS, RSA, UC, WTU). 3 mi E of Dos Cabezas, Maguire 11152 (DAO, GH, NY, UC, WTU). Mes al (ca. 7 mi W of Benson), Thornber 4312 ( ARIZ, OKLA, SMU). Dragoon, Trogstadt 1068 CARIZ. NMC). Ca. 3 mi E of Cochise Stronghold, Dragoon Mts., Towner 161 (D Gila Co.: 1 mi N of Black R., San Carlos Indian Reservation, Goodman 2 Hitchcock 1287 uer F, MO, NY, PH, POM, UC). 1 mi N of Blackriver Road, Granfelt in 1960 (ARIZ). Betwe n Globe and Cooley (Coolidge? ), Nelson 10372 (DS, MO, NY, mixture with C. * Pima Co.: Redington, Goodding in 1935 (ARIZ). Tucson Redington Road, San Pedro Valley, Brass 14282a (GH, NY). Pinal Co.: Near Oracle, ata Catalina Mts., Lewis 1079 (RSA, UC). Peppersauce Canyon, Santa Catalina Mts., 1 in 1937 (NY). Hills near Oracle, Harrison & Kearney 6673 (US). 7.7 to 7.9 mi SE of Oracle, iie T alina Mts., Towner 1, 3 (DS). Santa Cruz Co.: Mustang Mts., Pringle in 1884 (F, GH, , NY, POM, US). Near Sonoita, Harrison & Kearney 5713 (ARIZ, US). Sonoita to Elgin, on s O Fulton 11485 ( ARIZ, US). 7.5 mi E of Sonoita, Gregory 404, 405 (DAO, RSA, UC). 7.5 mi SE of Sonoita on road to Canelo, Towner 105 (DS). MEXICO. COAHUILA: Santa Rosa Mts., Marsh 1338, 1491 (F, OKLA, SMU, TEX). 27 mi E of Boquillas, Henrickson 11611b (TEX). 64 mi W of Cuatro Cienegas, Henrickson 7861 (TEX). 4.5 km E of Matrimonio Viejo, Johnston 10895 (TEX A s E, ~ Most abundant of the races of this species, ee hartwegii subsp. pubescens occurs widely in Texas and neighboring states. In central Texas, it is the only member of sect. Salpingia. Spreading trichomes and broad truncate— based leaves are strongly correlated in this form, and are characteristic of the central Texas populations and most others which are not affected by introgres- sion. The type of Oenothera greggii is tentatively included here, although the plants are stunted and somewhat lacking in distinctive characters. They are not typical of C. hartwegii subsp. pubescens and may represent hybrids or intro- gressants with subsp. hartwegii, which also occurs in that region. For these rea- sons, I have followed Shinners’ (1964) decisions and taken up the epithet “pubescens” for this taxon. 1977] TOWNER--CALYLOPHUS 87 Calylophus hartwegii subsp. pubescens is one of the most variable taxa in Calylophus, much of this perhaps stemming from the influence of introgression. The size of the flowers and of leaves and other vegetative parts all vary widely. Leaf margins may or may not be crinkled. Pubescence may consist wholly of spreading hairs or of these combined with shorter glandular or nonglandular pubescence. These characters appear to vary in response to genetic exchange with subspp. filifolius and hartwegii. Records of pollinators reported by Gregory (1964; as Oenothera greggii) in Terrell Co., Texas and Cochise Co., Arizona, included hawkmoths in the evening at both sites ( Hyles lineata, Manduca quinquemaculata, Sphinx dolli) and bees in the morning at the Cochise Co. site ( Megachile, Melissodes, Bombus). On a different date at the same locality in Arizona, I observed numerous halictid bees (Dialictus, Agapostemon, Evylaeus) and some bumblebees (Bombus) gathering pollen in the late afternoon. In the evening, hawkmoth ( Hyles lineata and Man- duca quinquemaculata) visitation was frequent. My field and greenhouse ob- servations showed a range in anthesis times extending from 2 hours before sun- set to about sunset. Thus flowers in this subspecies may not always be open early enough for significant afternoon visitation by bees. Ultraviolet-absorbing regions on the petals were found to be of small to moderate size. Each o four plants that were artificially pollinated was found to be self-incompatible. Tetraploidy and translocation heterozygosity are present in C. hartwegii subsp. pubescens, with tetraploidy occurring in one (Pecos Co., Texas, Munz & Gregory 23405) of the 18 populations which have been examined. A population inter- mediate between subsp. pubescens and subsp. hartwegii (Brewster Co., Texas, Munz Ç Gregory 23401) was also found to be tetraploid by Kurabayashi et al. (1962). Interchange heterozygotes comprised 15 of 20 plants examined for mei- otic configurations, occurring in 11 of 15 populations. An average of 1.2 trans- location heterozygosities per plant was calculated for this form, with a maximum of 3, the most frequent number being 1. No extra diminutive chromosomes have yet been observed in C. hartwegii subsp. pubescens. Configurations obtained from hybrids with other members of sect. Salpingia exhibited 0-2 translocation heterozygosities. 2 Most instances of introgression have been discussed in earlier sections. Two possible cases of hybridization with C. lavandulifolius were discovered for this taxon, perhaps the only subspecies of C. hartwegii which hybridizes in nature with C. lavandulifolius. One was a sight record from Guadalupe Co. New Mexico, where some plants with slight resemblances to C. lavandulifolius were found in two populations of C. hartwegii subsp. pubescens. The second was a single plant intermediate between the same taxa [between Globe and Cooley (?), Arizona, Nelson 10372, (RM)] observed in a mixed collection of the two typical forms. Numerous cases of sympatry without hybridization were observed, especially with C. serrulatus in Oklahoma, New Mexico, and the northern Texas Panhandle, and with C. berlandieri in central Texas and the Panhandle. Other instances included populations mixed with C. lavandulifolius in Culberson Co., Texas and with C. tubicula in Chaves Co., New Mexico. 88 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 2. Calylophus lavandulifolius (Torr. & A. Gray) Raven, Brittonia 16: 286. 1964 Oenothera lavandulaefolia 'Torr. & A. Gray, Fl. N. Amer. 1: 501. 1840. O. hartwegii Benth. var. lavandulaefolia (Torr. & A. Gray) S. Wats., Proc. Amer. Acad. Arts 8: 590. 1873. Galpinsia lavandulaefolia (Torr. & A. Gray) Small, Fl. S.E. U.S. 845, 1335. 1903. Oenothera hartwegi var. fendleri (A. Gray) A. Gray subvar. lavandulaefolia ( Torr. : A. Gray) H. Lév., Monogr. Onot 8. O. 'lavandulaejolia var. typica Mun Amer. J. Bot. 16: 704. 1929. Calylophus ‘hartwegii ( Benth.) Raven var. iavandulaejolius ray) Shinners, Sida 1: 345. 1964. v dd lavandulifolia Torr. " lavandulifolia; vedi N. Amer. Fl. ser. 2^ 5: 138. 1965. Calylophus oo osa lavandulifolius (Torr. & A. p ay) Towner & Raven, M: cay 20: 243. Oenothera lavandulaefolia Torr. & Gray var. glandulosa Munz, Amer. J. as 15 705. 1929. Galpinsia lavandulaefolia (Torr. & A. Gray) Small var. glandulosa (Munz) Moldenke, Phytologia 2: 134. 1946. rype: United States, Nevada, White Pine Co., Ely, 30 July 1923, M. E. Jones (POM). Similar to Calylophus hartwegii. Suffrutescent perennial from a stout woody caudex, caespitose, sometimes appearing more or less tufted; stems several to many, moderately branched, spreading-decumbent to more or less ascending, 0.4-2(-3) dm high; plant densely gray-strigulose throughout. Leaves dense on the stem, sessile, usually ascending, linear to narrowly lanceolate or narrowly ob- lanceolate, 6-50 mm long, 0.8-6 mm wide, the tip acute or obtuse, the base acute- attenuate, the margin entire or nearly so, occasionally slightly undulate, in- frequently revolute; small axillary leaves present, 2-10 mm long; lowest stem leaves somewhat wider and more oblanceolate than above. Floral tube 25-60 mm long, 5-15 mm wide at the throat, minutely strigulose or glandular-pubescent without, sometimes with purple longitudinal lines and base, occasionally fading pinkish upon wilting. Sepals 8-20 mm long, 3-8 mm wide, with free tips 0.3-3 mm long, usually with purple marginal stripes. Petals 12-28 mm long, similar in width, usually fading pinkish to purplish, highly ultraviolet-reflective, with a small basal ultraviolet-absorptive spot, rarely medium-sized. Filaments 6-12 mm long; anthers 5-11 mm long. Style 30-75 mm long, glabrous above, minutely pubescent below; stigma 2-5 mm broad; ovary 4-16 mm long, 1-2 mm wide. Capsule 6-25 mm long, 1-3 mm wide; seeds 1.5-2.5 mm long. Self-incompatible. Gametic chromosome number, n — 7 TYPE: UNITED States. On plains, probably along the South Platte River in southwestern Nebraska or northeastern Colorado, June or July 1820, Edwin James (PH). The approximate locality was taken from McKelvey (1955: 213- 219) and is at the eastern limit of the range of this species. Distribution (Fig. 17): Local and sparse, on sandy and rocky, often cal- careous soil, on high plains and in mountains, frequently with Juniperus, Pinus monophylla or Pinus edulis, Cercocarpus, Artemisia tridentata, occasionally in lower zones with Larrea divaricata or in higher zones with Pinus ponderosa, from southern Fall River Co., South Dakota, southeastern Wyoming, and far western Nebraska, through western Kansas, Colorado, eastern and southern Utah, north- western Oklahoma, and the Texas Panhandle to Trans-Pecos Texas, central . Nuevo León, central New Mexico, central Arizona, and east-central and southern Nevada. Occurring from elevations of ca. 600 m (2 mi W of Hays, Ellis Co., 1977] TOWNER—CALYLOPHUS 89 SURE 17. Distributions of Calylophus toumeyi (triangles), C. 8 (dots), C. tubicula subsp. awama (open circles), and C. tubicula subsp. strigulosus (squar 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Kansas) to ca. 2,750 m (Lee Canyon, Spring Mts., Clark Co., Nevada). Flowers April to August. Representative specimens examined: INrTED STATES. SOUTH DAKOTA: Fall River Co.: Rocky dry ridges, Over 18074 (RM). WYOMING: Goshen € 5o.: 4 mi N of La Grange, Stephens & Brooks 22903 (DS). Laramie Co.: Hartville, Nelson 8328 (GH, MO, NY, US). 20 mi W of Pine Bluffs, Porter & Porter 8153 (DS, POM, UC). SE edge of Pine Bluffs, Stephens & Brooks 22873 (DS). Platte Co.: Whalen Canyon, Nelson 526 (GH, MO, NY, US). Near Guernsey, Porter 3557 (DS, SMU, TEX, UC, TL NEBRASKA: Box Butte Co.: 5 mi E and 5 N of Hemingford, Stephens & Brooks 24542 (KANU). Chase Co.: 10 mi N of Imperial, Brown 1255 (NEB). Garden Co.: 2 mi S of Lewellen, Stephens & Brooks 11519 (KANU). Morrill Co.: Angora, Pool in 1912 (MO, NEB). 4 mi N of Broadwater, Stephens & Brooks 13907 (DS, KANU). Scott's Bluff Co.: 1.5 mi W and 5 S of Melbeta, Stephens 5484 (KANU). coLorapo: Baca Co.: 9 mi S and 2 E of Walsh, Ste 915 ns & Brooks 21820 (DS). Bent Co.: 15 mi SE of Las Animas, Stephens & Brooks 22003 (DS). 4.5 mi W of Prowers, Stephens & Brooks 21989 (DS). Dolores Co.: Just W of Nor ihdale: Anderson 3138 (DS). Huerfano Co.: 19 mi NE of Walsenburg, Stephens & Brooks 22229 (DS). Kit Carson Co.: 5 mi E of Flagler, Stephens & Brooks 22633 (DS). Las Animas Co.: 6 mi N and 4 E of Andrix, Stephens & Brooks 21900 (DS). Otero Co.: 2 mi S of Manzanola, Stephens & Brooks 22315 (DS). Sedgwick Co.: 1 mi S of Julesburg, Stephens & Brooks 24060 (KANU). Kansas: Clark Co.: 8 mi N 7 DR Horr E248 (COLO, DAO, F, GH, KANU, LL, OKL, RM, SMU, TEX, UC). Ellis 2 mi W of Havs, Bondy 77 (ARIZ, F, MO, NMC, OKL, PH, RM, n Gove Co.: 20 y 8 ud 3 E of Oakley, Lathrop 3374 (KANU, SMU). Meade Co.: 12 mi E of Meade, Horr 3532 (KANU, TEX, US). Morton Co.: 7 mi N and 4 W of Elkhart, 0 8877 (KANU). scott Co.: Horsethief Canyon, Scott County State Park, Fearing & Latham in 1950 (GH, KANU). Trego Co.: 14 mi S of Ogallah, McGregor 17124 (KANU). OKLAHOMA: Beaver Co.: E edge of Elmwood, Stephens & Brooks 21745 (DS). Harper Co.: 10 mi S of Buffalo, Gooaman 2394 (MO, NY, OKL, POM, UC, WTU). Texas Co.: 5.5 mi E of Hardesty, Stephens & Brooks 21776 (DS). Woods Co.: Near pees Stevens 252 (DS, GH, NY, OKL, SMU). Texas: Brewster Co.: Foothills of Glass Mts., 7.7 mi NE of U.S. 90 on U.S. 67, Towner 29 (DS). Culberson Co.: 25.6 mi SW of White’s City. E side of G uadalupe Mts., Towner 24 (DS). Hudspeth Co.: 31 mi E of El Paso, Hueco Mts., Tharp 46071 (F MO. TEX). NEw Mexico: Colfax Co.: Near Raton, Nelson & Nelson 4681 (DS, RM, US). Eddy = Near Three Forks of Rocky Arroyo, Med Mts., Wilkens 1711 e pod Co.: Hell Canyon, Magdalena Mts., Herrick 274 (US). Tominés Co.: 3.0 mi NE of Duran, D 19129 (DS). 5.8 mi SW of Duran, Raven 19133 (DS). Ca. 7.5 mi W of y eee on U.S. 60, Towner 120 (DS). ARIZONA: Coconino Co.: Rim of Canyon Diablo, Two Guns, Demaree 44216 (ARIZ, PH, RSA, SMU). E rim of Canyon Diablo, 'Two Guns, Towner 114 (DS). 10 mi SE of Tuba City, Peebles 13363 (GH, US). 10.9 mi S of Bitter Springs, Mosquin & Mosquin 4247 (DS). Mojave Co.: Ca. l mi from rim of canyon, Toroweap Valley, McClintock 52-512 (ARIZ, NY). Navajo Co.: 45.0 mi NW of Concho, Towner 115 (DS). UTAH: ange Co.: Juniper zone, below Moon Lake, Graham 6412 (MO, POM). Emery Co.: 50 mi N of Hanksville, San Rafael Swell, m 9201 (DAO, DS, NY, POM, WTU io Garfield 000. Red Canyon, 100 mi W of Bryce on, Preece 2480 (C OLO, POM, SMU). Bryce Canyon, Goodman d . Ne 1566 (DS, Ca CH. POM, RM, UC). 10 mi E of Escalante, Holmer ren & Nielsen 7734 (DS, POM, RM, UC, WTU). Millard Co.: Tunnel Springs, Desert Game Range, 3 8553 (ARIZ, POM). San Juan Co.: Tuwa Canyon, Natural 5 1 Monument, Welsh & Moore 2294 (NY). Uinta Co.: Willow Creek, S of Ouray, Holmgren 1882 (KANU, WTU). Washington Co.: 10 mi N of the Beaver Dam summit of U.S. 91 and 5 mi NW of the highway, Wiens 3917 (WTU). NEVADA: Clark Co.: Old Saw Mill site, Sheep Mts., 6,600 ft, Alexander d» Kellogg 1757 (GH, UC, US, WTU). Rocky ridge S of Deer Creek, Charleston (Spring) Mts., 2,670 m, Clokey & Clokey 7605 ( ARIZ, CAN, D O, DAO, DS, F, GH, KANU, MO, OKL, PH, POM, RM, RSA, SMU, TEX, UC, US, WTU). 4.8 mi N of Kyle Canyon on road to Lee Canvon, Charleston Mts., Towner 101 (DS). Lee Canyon, 0.7 mi W of jct. to Kyle Canyon, Charleston „ Towner 104 (DS). es Co.: Panaca Valley and vicinity, Gentry 131 (ARIZ, DS, 12 US). White Pine Co.: 3 mi S of Ruth, Moore 346 (DS, POM). 5.1 mi S of U.S. 50, on eastern road to Hamilton, opus o Solbrig 13550 (DS). 2 mi w of Ely, an 2897 (KSC). EXICO. NUEVO LEÓ 2 E e S of San Roberto Junction on Mexico 57, Sanderson 291, 292 ( TEX); Turner 6357 (T . 16 mi S of San Roberto Junction, Reveal et al. 2652 (DS). Near summit of N.L. Am ay ron W of Galeana, d 288, in part ( TEX). 1977] TOWNER—CALYLOPHUS 91 The limits of this species are approximately those given by Munz (1929). Some collections included here by Munz clearly belong with Calylophus hart- wegii subsp. hartwegii, e.g., Zacatecas, gravelly soil, Purpus in 1903 (UC). The two taxa are easily confused but differ in the shorter sepal tips, d pubes- cence, and leaves which are usually broader and obtuse-tipped in C. lavanduli- folius. A species of broad distribution, C. lavandulifolius occurs for the most part to the north and northwest of the range of C. hartwegii. Isolated collections have been made, however, throughout much of the range of the latter. Where C. lavandulifolius occurs with C. hartwegii, it tends to occupy higher elevations than any race of that species, except for C. hartwegii subsp. fendleri. Plants of C. lavandulifolius are typically slow-growing, small, and sparsely distributed, and are relatively inconspicuous when mixed with populations of the more abundant C. hartwegii. Hybridization with other species of Calylophus apparently occurs only rarely. “vidence for gene exchange stems only from the two collections mentioned previously which were intermediate between C. lavandulifolius and C. hartwegii subsp. pubescens. Instances of populations of C. lavandulifolius contiguous with those of other taxa have also been mentioned above, and include contact with C. tubicula subsp. tubicula, C. tubicula subsp. strigulosus, C. hartwegii subsp. pubescens, C. hartwegii subsp. fendleri, and C. serrulatus. Variation within C. lavandulifolius is not clearly correlated with geography, and division into subspecies seems inadvisable. The glandular pubescence of the floral tube and calyx used by Munz to distinguish Oenothera lavandulaefolia var. glandulosa does not appear to vary in any meaningful pattern or in associa- tion with any other character. Considerable variation within the species occurs in leaf dimensions, leaf margin (undulate, revolute, or plane), width of the floral tube, and in other floral characters. Visitors to a population in Clark Co., Nevada (Towner 195) consisted en- tirely of hawkmoths ( Hyles lineata, Manduca), which were active between dusk and dark. Anthophorid and halictid ( Agapostemon) bees were seen on flowers 2 hours before sunset at a population in Torrance Co., New Mexico (Towner 120). Anthesis in the field and greenhouse ranged from 3 hours before sunset to sunset. In the population in Clark Co., Nevada the median time of anthesis was 1% hours before sunset. Ultraviolet absorption patterns on the petals tend to be small in this species. The foregoing facts suggest that hawkmoth pollination predominates in C. lavandulifolius, but that bees may also play a role at certain localities. Two plants were self-pollinated and found to be self-sterile. Cytological variation in Calylophus lavandulifolius occurs in the form of translocations and extra diminutive chromosomes. Seven of 10 plants from 5 of 6 populations were interchange heterozygotes. An average of 1.2 heterozygosities per plant was calculated. Three individuals from separate populations had extra chromosomes, which consisted of 1 or 2 diminutive pairs. Configurations dis- played by hybrids of C. lavandulifolius with other taxa in sect. Salpingia showed 2-3 translocation differences and occasional inversion differences between the 99 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 64 parents, a greater cytological divergence than shown by other crosses within the section, 3. Calylophus toumeyi (Small) Towner, comb. nov.—Fics. I, 3. Galpinsia toumeyi Small, Bull. Torrey Bot. Club 25: 317. 1898. Oenothera hartwegii Benth. var. toumeyi (Small) Munz, Amer. J. Bot. 16: 708. 1929. O. toumeyi (Small) Tidestrom, Proc. Biol. Soc. "Wash. 48: Al. 1935. Calylophus hartwegii ( Benth aven var. toumeyi (Small) Shinners, Sida 1: 341. 1964. C. hartwegii subsp. toumeyi (Small) Towner & Raven, Madrono 20: 243. 1970 =: Similar to Calylophus hartwegii. Suffrutescent perennial from a stout woody caudex; stems several, sparingly branched or unbranched above, ascending to erect, 1.5-6(+) dm high; plant subglabrous to minutely strigulose throughout. Leaves sparsely distributed on the stem, sessile, more or less spreading, narrowly lanceolate, 10-35 mm long, 1-7 mm wide, the tip acute, the base acute-attenuate, the margin entire to obscurely and sparsely serrulate, not undulate; conspicuous fascicles of small leaves 2-25 mm long in nonfloriferous axils; lowest stem leaves usually tending towards oblanceolate shape. Floral tube (15-)30-60(-70) mm long, 5-14 mm wide at the throat, yellowish, fading orangish to brick red upon wilting. Sepals 10-25 mm long, 3.5-6 mm wide, with free tips 2-9(-12) mm long, colored as the floral tube. Petals 10-20 mm long, similar in width, intensely lemon yellow, fading orangish to brick red, moderately ultraviolet-absorptive throughout. Filaments 4-12 mm long; anthers 6-10 mm long. Style 35-70(—-80) mm long, glabrous above, minutely pubescent below; stigma discoid to squarish, 1.5-4 mm broad; ovary 6-20 mm long, 1-2 mm wide. Capsule 10-50 mm long, 1.5-4 mm wide, thin walled, sometimes almost papery, dehiscent only in the dis- tal half; seeds 2-3 mm long. Self-incompatible. Gametic chromosome number, n=7 TYPE: UNITED STATES. ARIZONA: Cochise Co., Chiricahua Mountains, 25 July 1894, J. W. Toumey 197 (NY); Munz, Amer. J. Bot. 16: 708. 1929. Distribution (Fig. 17): Local and uncommon on shaded rocky slopes or disturbed areas in pine-oak forest, from the Santa Rita, Huachuca, and Chiricahua mts. in Santa Cruz and Cochise cos., Arizona, and the Mogollon Mts. in southern Catron Co., New Mexico, south through northeastern Sonora in the Sierra Madre Occidental to west-central Chihuahua. From elevations of ca. 1,500 m (Stone Cabin Canyon, Huachuca Mts., Santa Cruz Co., Arizona) to 2,600 m (summit of the José Mts., Sonora). Flowers mostly from July to October, but some popu- lations in Mexico as early as May. Representative specimens examined: UNITED STATES. NEW MEXICO: Catron Co.: On or near the West Fork of the Gila R., Mogollon us ` Metcalfe 555 ( ARIZ, GH, MO. NMC, US). arizona: Cochise Co.: Huachuca Ats., Harrison & Kearney 5773 (US). Fort a Wilcox in 1892 (NY). Near Fort Huachuca, “Wilcox 253 (US). Huachuca Mts., 7000 ft, Jones in 1903 (DS, POM, US). Huachuca Mts., Toumey in 1894 (GH, RM US). lm er's Canyon, Huachuca Mts., Gilbert in 1892 (NY). Tanners Canyon, Huachuca Mts., emmon in 1882 (UC). Ramsey's Canyon, Huachuca Mts., Goodding 786 (RM, US). Miller Canyon, Huachuca Mts., Carter in 1936 (NMC). Carr Peak, Huachuca Mts., 6500 ft, Benson 10500 ( ae Carr Peak, Huachuca Mts., Goodding 222 (ARIZ, GH, NEB, NY, OKLA, RM). 5.6 mi up Carr r Canyon road from Arizona 92, Huachuca Mts., Towner 106 (DS). Reef Mine, Huachuca Mts., 7100 ft, Gould 1977] TOWNER—CALYLOPHUS 93 1475 (ARIZ, UC). Garden Canyon, Huachuca Mts., Harrison & Kearney 5773 (ARIZ). Near Fort Huachuca, Huachuca Mts., Lemmon 2700 (GH). Rucker Canyon, upper left fork, Chiricahua Mts., Blumer 2025 (F). Sugar Loaf Mt., Chiricahua Mts., Darrow in 1937 (ARIZ). N slope of Sugar Loaf, Chiricahua National sil ie bis $380 (ARIZ). Sugarloaf Tras just below tunnel, Chiricahua National Monument, Towner 107 (DS). Near summit of pass, Chiricahua Mts., Goodding 165-47 (ARIZ). Pine 3 Ghirieahna Mts., 6700 ft, Blumer 1610 (ARIZ, DS, F, MO, NEB, e NY, RM, US). Ida Peak, along Telephone Trail, Chiricahua Mts., 8,000 ft, Stone 517 (P I. RM). Pinery Canyon, Chiricahua Mts., 7,000 ft, Barr 64-353 (ARIZ). 12.4 mi W of jet. of — 186 & 181, in Pinery Canyon, Chiricahua Mts., ca. 7,000 ft, Towner 164 (DS). 12.5 mi W of jet. Arizona 186 & 181, Towner 171. 1 mi below Onion Saddle, E. side of Chiricahua Mts., Kaiser 49-209 (ARIZ). Crest Trail, Chiricahua Mts., 7,000 ft, Fa r 136 (ARIZ). Gut Saw Canyon, Chiricahua Mts., Goodding 2339 UC). Wonderland of Rocks, Chiricahua Mts., Darrow in 1937 (GH, NY E Bonita Canyon, Chiricahua Mts., Henderson in 1933 (TEX). Outlaw Canyon, Wi. sis Mts., Goodding 2339 (RM). Pima Co. (?): Sabino Canyon, Catalina Mts., 3,000 ft, Jones in 1903 (MO, specimen immature; either not a this species or locality veia d Santa Cruz Co.: Madera Canyon, Santa Rita Mts., 5,900 ft, Darrow 2614 (ARIZ). Madera Canyon, Santa Rita Mts., Peebles et dl. 4545 (A RIZ, US). Stone Cabin Canyon, Santa Rita Mts., 5,000 ft, Thornber in 1903 (ARIZ). Santa Rita Mts., 6,000-8,000 ft, Pringle in 1881 (F GH). Santa Rita Mts 7,000 ft, Darrow & Arnold in 1936 (MO, OKL). Upper Madera en Santa Hita Mis. Clark 1 12351 (GH, OKL). Along trail I: Mount Wrightson to White House Canyon, Santa Rita Mts., 7,000 ft, Parker et al. 5835 (ARIZ, NY ! EXICO. sONORA: Puerto de los Aseríados, region of the Río de Bavispe, White 3190 (ARIZ). Summit mi Bar José Mts., 8,600 ft, Mearns 1606 (DS, US). Cananea, Murdoch in 1914 (V). CHIHUA 48 mi W of Matachic on road to Ocampo, 8,400 ft, Wiens 3445 ( COLO, DS). Mojarachic (Maguarachie? ), Knobloch 5094 (F). Mts. NW of C Chihuahua. Le iu ur in 1936 (MO, TEX, UC, US). San José de Pinal, Río Mayo, 7,000 ft, Gentry 2587 ( ARIZ, MO, POM, UC, US). Mts. SW of Chuchuichupa, Hartman 712 (F, NY, UC, US). 1 8 5 NW railroad. km 85, Barlow in 1911 (F). Rice 10 tween Rio Chico and Río Caballo, Mexican NW railroad, d Divide, Barlow in 1911 (F). Carretas, White 993 (ARIZ). 130 mi W of Chihuahua Citv, 8,500 ft, Russell in iom (UC). Canon boat 10 mi SE of Madera, Muller 3428 (UC i “Gosyaninë. rh yon, Sierra Madre, Jones in 1903 (POM). Salto de Babicora, Le Sueur 1407 (F). W from Pen (now Juan Mata Ortiz), Sierra Madre, Barlow in 1911 (F). Sierra Madre, Nelson 6088 (US). No specific locality, Le Sueur 101 (F, TEX). 7 This most distinct of the large-flowered members of sect. Salpingia is readily separated using any of several characters, including its exclusively montane dis- tribution, fascicles of large axillary leaves, tall erect stems, long sepal tips, partially dehiscent capsule, and unusual ultraviolet absorption pattern on the petals. Apparently completely allopatric to the other taxa of sect. Salpingia, Calylophus toumeyi experiences no current genetic exchange with them. The absence of any intermediate collections and the number of characters which show discontinuities from C. hartwegii indicate the validity of specific status for C. toumeyi, which was not recognized in an earlier publication (Towner & Raven, ) Distributed in the mountains of southeastern Arizona, southwestern New Mexico, and northeastern Mexico, this species is physically and ecologically separated from other members of sect. Salpingia. The blooming period is un- usual for the genus, occurring in late summer and early fall in response to sum- mer rainfall. Earlier flowering is prevented by the late, dry spring, characteristic of montane areas in this region. Flower visitation to C. toumeyi seems sporadic, as insects were seen only once in significant numbers during several attempted studies. On that occasion, in the Chiricahua Mts., of Arizona (Towner 238), bees of the genus Lasioglossum were active gathering pollen shortly before dusk, and again after sunrise. Hawk- 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 moths were abundant visitors in the evening. Morphological characters and anthesis times suggest that hawkmoths are the principal pollen vectors. The floral tube attains a length of 70 mm in some specimens, the maximum seen in Calylophus, and is perhaps a response to pollination by the genera of sphingids with longer tongues (cf. Gregory, 1964). Moderately absorptive to ultraviolet light over their entire area, the petals have no contrast pattern, nor do they contrast with vegetative parts (Fig. 12). Anthesis occurs % to 1% hours before sunset. Bees are therefore not likely to be regular and significant contributors to pollination. Self-incompatibility is probably typical for this species. Only one plant was checked by self-pollination, and it was self-sterile. In one field study, a popula- tion was observed to have set no seed in spite of having been in flower for several weeks. The same population showed over 60% fertile capsules on the date of the cited pollination study. Three plants from two Arizona populations were examined cytologically and proved to be heterozygous for two translocations apiece. Multivalent associa- tions included a ring of 6 chromosomes in 1 plant and 2 rings of 4 chromosomes in the 2 others. One of each type of configuration was found in a population from the Chiricahua Mountains (Towner 107), indicating that at least 3 trans- location polymorphisms were present in the population. Chromosome determina- tions from hybrids were not obtained because of the difficulty of crossing C. toumeyi and C. hartwegii and because of the scarcity of floral buds on those hybrids which were produced. 4. Calylophus tubicula (A. Gray) Raven, Brittonia 16: 286. 1964. Oenothera tubicula A. Gray, Pl. Wright. 1: 71. 1852. Herbaceous or slightly suffrutescent short-lived perennial, arising from a slender woody caudex; stems one to several, sparingly branched above, sub- decumbent-ascending to nearly erect, 0.4—5.3 dm high; plant with short glandular pubescence throughout, or with some parts minutely strigulose. Leaves = dense, subsessile, ascending, linear to ovate or obovate, 7-46 mm long, 0.7-11 mm wide, the tip acute, sometimes obtuse in lowermost leaves, the base acute-attenuate, the margin entire or sparsely and shallowly serrulate, occasionally slightly un- dulate; fascicles of small leaves 2-15 mm long in nonfloriferous axils; lowest stem leaves more frequently oblanceolate than above. Inflorescence dense, with buds crowded near the stem apex; buds terete. Floral tube funnelform in upper one- half or more, often tubular below, 5-25(-33) mm long, 3-10 mm wide at the throat in pressed specimens, the inner surface glabrous above to densely pubes- cent basally, yellow, sometimes fading pink, and more rarely, purplish. Sepals 3-13 mm long, 2-6 mm wide, with subulate free tips 0.5-2 mm long, plane, yellow, rarely with purple marginal stripes, only infrequently fading pink or purplish. Petals suborbicular to obovate-truncate, 5-20(-25) mm long, similar in width, infrequently fading pink to purplish, highly ultraviolet-reflective, with a large basal ultraviolet-absorptive spot. Stamens subequal; filaments 1-6 mm long, glabrous to minutely pubescent; anthers 2-7 mm long, sparsely and minutely 1977] TOWNER—CALYLOPHUS 95 . pubescent. Style 9-30(—40) mm long, usually exceeding the stamens, glabrous above, minutely pubescent below; stigma discoid to squarish, 1-2.5 mm broad; ovary 4-11 mm long, 0.5-1.5 mm wide. Capsule 8-19 mm long, 1.5-2.5 mm wide, moderately thin walled, completely dehiscent; seeds 1.0-1.4 mm long, obovoid, angled, truncate at the apex. Self-incompatible. Gametic chromosome number, =7. TYPE: UNITED STATES. TEXAS: prairies beyond the Pecos River, probably in eastern Pecos Co., August 1849, Charles Wright 197 (in part = 821; GH). Distribution (Fig. 17): Primarily on limestone soils in arid lowlands, but occasionally in montane areas, from Guadalupe Co., New Mexico, south to west- ern Texas, thence northeast to Howard Co., Texas and south to northern Zacate- cas, south-central Nuevo León, and southwestern Tamaulipas. From ca. 600— 1,800 m elevation. Flowers April to August. Cytological relationships, interfertility, and a number of morphological char- acters demonstrate a close affinity between Calylophus tubicula and the other members of sect. Salpingia. However, the short-tubed, funnelform flowers (Fig. 4) and morning anthesis of this species constitute a phenetic similarity to sect. Calylophus, a relationship likely due to evolutionary convergence. Calylophus tubicula is distinguished from its closest relatives by those same characters. In addition, individuals of this species are shorter-lived than in other forms in the section. The plants rarely have large taproots, and are difficult to maintain for more than one or two years in cultivation. Self-sterility was found in six plants from two populations of C. tubicula subsp. tubicula, and no evidence of self-compatibility was seen in any cultivated or wild plants of either subspecies. Stigmas were invariably well exserted and no greenhouse specimens of either subspecies ever set seed spontaneously. An- thesis occurred just before dawn (% to 1% hours before sunrise) at two colonies of C. tubicula subsp. tubicula in Eddy Co., New Mexico and in cultivated rep- resentatives of both subspecies. Stamens, the stigma, and a large spot at the base of each petal are ultraviolet-absorptive, contrasting markedly with the rest of the petal surface, which is highly reflective. Flower visitors at field sites (Towner 14, 15) of C. tubicula subsp. tubicula consisted primarily of small halictid bees of several genera, especially Evylaeus, Dialictus, and Agapostemon. These were most active from just before sunrise to mid-morning, gathering both pollen and nectar. Infrequent visits by hawkmoths (Hyles lineata) were observed shortly before sunrise. In some colonies, removal of pollen was virtually complete by afternoon. Field data, anthesis times, and flower morphology indicate that this species is predominantly bee-pollinated. A strong inference can be made that C. tubicula evolved from a moth-pollinated ancestor resembling C. hartwegii. This is based on the relatively long floral tubes in some C. tubicula and on the close phenetic and cytogenetic relationship of the two species. Relative to other forms of Calylophus, C. tubicula has a low level of chromo- some heterozygosity. Eight of 14 plants examined had multivalents, and no evidence of inversions, diminutive chromosomes, or polyploidy was found. The 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 average number of observable translocation heterozygosities per plant was 0.7, as compared with 1.0 for the rest of sect. Salpingia and 1.9 for C. berlandieri. Individuals from certain populations of C. tubicula may approach or exceed the minimum floral tube length seen in C. hartwegii. In such cases, C. tubicula generally retains the wider funnelform shape of the tube. Other populations show vegetative characteristics which suggest recent introgression with or deriva- tion from C. hartwegii. These cases will be discussed under the subspecies. Differences of pubescence, leaf shape, floral pigments, and ecological dis- tribution distinguish the two subspecies listed below. The extremes of each form are quite distinct, but individuals from several collections in Mexico cannot be assigned with certainty to either taxon. KEY TO SUBSPECIES a, Glandular-pubescent oup leaves narrowly lanceolate to ovate; flowers rarely fading reddish or purple 4a. EE tubicula aa. Minutely grey-strigulose on the ov vary and upper stems; leaves linear to narr lanceolate; flowers commonly fading reddish or purple 4b. subsp. 8 4a. Calylophus tubicula (A. Gray) Raven subsp. tubicula— Fi. 4. isi ce tubicula (A. Gray) Small, Bull. Torrey Bot. Club 23: 186. 1896. Oenothera artwegi Benth. var. tubicula (A. Gray) H. a Monogr. Onoth. 335. ; Oenothera tubicula A. Gray var. ee A. Gray, Pl. Wright. 1: 71. 1852. Type: United e Culberson Co., on the VERONA Mts., October 1849, Charles Wright 197, in 1 = 13380 (GH, holotype; US, 5 The collection probably came from Texas and not New adis (McKelvey, 1955: Oenothera x serrulatoides H. Lév., oe Onoth. 335. 8. rype: United States, Texas, Co., valley of the Pecos and towards the asnaq ae 1851, septa 1 1077 (MO, holotype; GH, NY, PH, isotypes). Remark on type s sheet: ais hybride de bicula X serrulata?” Galpinsia ETENN A. Nels., . J. Bot. 21: 575. 1934. rype: United States, New exico, Eddy Co., a 11 vicinity of Carlsbad Caverns, May 1930, Gladys Convis 36 (RM). Published 5 Galpinsia carlsbadia Nels., ree d. Bot. 23: 269. 1936. TYPE: United States, New exico, Eddy Co., r the Caverns, Carlsbad National Park, 24 May 1931, Aven Nelson 11396 (RM, 9 8 "DS. NY, POM, isotypes). With short glandular pubescence throughout. Leaves narrowly lanceolate to ovate or obovate, 7-46 mm long, 0.7-11 mm wide, usually entire or nearly so. Flowers rarely fading reddish or purplish upon wilting. Self-incompatible. Gametic chromosome number, n = 7. Distribution (Fig. 17): Colonial, primarily on limestone soils, in flat arid grasslands, often with Larrea divaricata and Yucca, from Guadalupe Co., New Mexico, south in the western side of the Pecos River drainage to western Texas, where occurring from Culberson Co. east to Howard Co., thence south through Presidio, Brewster, and Terrell cos., and probably most of central Coahuila, to northern Zacatecas, southwestern Nuevo León, and southwestern Tamaulipas. Elevational distribution from ca. 600 m (10 mi E of Dryden, Terrell Co., Texas ) to ca. 1,400 m (between Santa Rosa and Vaughn, Guadalupe Co., New Mexico). Flowers April to August. 1977] TOWNER—CALYLOPHUS 97 Representative specimens examined: NITED STATES. NEW MEXICO: Chaves Co.: Ca. 4 mi N of Roswell, Towner 127 (DS). 3 Co.: 4 mi W of Hope, Munz & Gregory 23350 (RSA). Memorial Hospital, N end of Carlsbad, [ey & Gregory 23353, 23354, 23. 356 ert UC). 0.6 mi W of Hope, Totner 14 (DS). L7 mi NE of Ho ope, Towner 16 (DS). 4.5 mi S of Carlsbad, Towner 17 (progeny only, DS). 7.6 mi NE of White's City, Towner 178 (DS). Otero Co.: Ca. 5 mi W of Elk (1 plant), Towner 108 (DS). Texas: Bre dou 41 mi S of Alpine: Anderson 3030 (DS). Flats near Old Blue, Glass Mts., W D Ws 1 (DS, POM, TEX). 15 mi E of Marathon, Munz & Gregory 23400 (RSA, UC). Ca. 10 mi E of Alpine, Sperry T1095 (UC, US). 6 mi S of Marathon, Rollins & Chambers 2766 (DS, GH, POM, RM, UC). Culberson Co.: 3 mi SW of New Mexico line on U.S. 180, Munz & Gregory 23360 (RSA). 9 mi E of Van Horn, Waterfall 4162 (ARIZ, GH, MO, NY). Ector Co.: W of Odessa, Lundell & Lundell 16921 (LL). Jeff Davis Co.: 8 mi S Siok Fort oo Munz Ü Gregory 22385 (RSA, UC). Pecos Co.: “Mesa s slope,” Tharp 43-731 (OKL, OKLA, RM, TEX, CC). Ca. 20 mi W of Sanderson, Warnock & McBryde 14904 (LL, TEX). n 1i E of Fort Stockton, Warnock 5164 (LL, SMU). 30 mi NE 5 Fort Stockton, Ownbey & Ownbey € (MO, POM, RM, RSA, UC). Presidio Co.: Up E between Long Draw and Capote Draw on road from ae to Ruidosa, e 1 SMU). Cleveland Ranch, near di Mts., Hinckley (GH 5 mi N of Marfa, 1 d Gregory 23389 (RSA, UC). Reeves Co.: On U.S. 80, 3 mi " of intersection with U.S. 290, Munz & Gregory 23370, 23371, 23372 (RSA, UC, WTU). U.S. 80, 1 mi E of intersection with U.S. 290, N edge of the Davis Mts., Waterfall in 1943 E O T (CH, MO, NY). Plains W of Pecos, Tracy & 8 155 144 (F, GH, MO, NEB, NY, ' Terrell Co.: 10 mi E of Dryden, Parks et 2 6 (TEX). Upton Qo: 20 mi SE of C 'ane, haven & Gregory 19240. Val Ve de d L Fisher 290 (US). Ward Co.: N of Pyote Lundell & Lundell 11379 (POM, SMU MEXICO. ZACATECAS: 18 km w of 2 oncepción del Oro, Stanford et al. 590 (DS, MO, NY). SAN LUIS POTOSI(?): “Prov. de San Luis,” Octoust 1050 (P). This subspecies occurs primarily on arid calcareous flats and outcrops southeastern New Mexico and western Texas. Its distribution in northern Mexico is very poorly known, and may include much of Coahuila in addition to the localities listed. In southern New Mexico, it occurs in the Pecos River Valley and on the plains to the west of it. Other taxa of Calylophus supplant C. tubicula on the higher plains to the east of the river (the “Llano Estacado” The broader leaves and glandular pubescence are the primary characters dif- ferentiating C. tubicula subspp. tubicula and strigulosus from one another. In- trogression between the two seems to occur in Nuevo León, T amaulipas, and Coahuila. Intermediate specimens are cited under subsp. strigulosus. Twelve plants from 9 populations have been examined during meiosis; 6 had 5 bivalents and a ring of 4 apiece, while 6 had 7 bivalents. Experimental hybrids were obtained with difficulty between C. tubicula subsp. tubicula and C. ber- landieri subsp. berlandieri. These plants were weak and essentially sterile (0-1% pollen fertility). They showed low chiasma frequency and gave evidence of at least 6 major translocation differences between the forms. Crosses of C. tubicula subsp. tubicula with other members of sect. Salpingia produced progeny which were structurally homozygous or heterozygous for 1 or 2 translocations. Such crosses produced good seed set and reasonably fertile hybrids, although the hy- brids frequently showed poor germination and viability. Introgression between this subspecies and C. hartwegii subsp. fendleri has been imputed by Munz (1965). Actually, considerable differences in elevational distribution separate C. hartwegii subsp. fendleri and C. tubicula subsp. tubicula, the former occurring from ca. 1,500 to 2,150 m elevation in New Mexico. This should make contact between these taxa infrequent. In the Carlsbad Caverns 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 area, individuals with broad leaves and long floral tubes have been treated as a distinct taxon, Galpinsia carlsbadiana, and as possible hybrids between the forms mentioned above. The latter possibility seems remote, since I have discovered no records of C. hartwegii subsp. fendleri within 160 km of the Caverns area. Taxo- nomic recognition is unwarranted also for the reason that the long-tubed forms occur together with typical C. tubicula in many populations throughout the range of subsp. tubicula. It is uncertain whether this pattern should be in- terpreted as a result of present or past genetic exchange or as spontaneous vari- ation within C. tubicula. Collections of C. tubicula subsp. tubicula showing long floral tubes include the following: 4 mi N of Carlsbad Caverns, Eddy Co., New Mexico, Porter & Porter 8986 (DS, RM; of all collections, this is the most hybrid- like, being very similar to C. hartwegii subsp. fendleri). 45 mi S of Pecos, Pecos Co., Texas, Moore & Moore 21 (NY, SMU, UC). 1-5 mi NW of Notrees, Ector Co., Texas, Collins 82 (OKLA). Borrow pits N of Pecos, Reeves Co., Texas, Nel- son & Nelson 4989 (DS, MO, RM, TEX, UC, US). In general, C. tubicula subsp. tubicula was not found growing together with other forms of Calylophus, perhaps because of its restriction to xeric sites at low elevations. In Chaves Co., New Mexico, two individuals of C. hartwegii subsp. pubescens ( Towner 126) were found in a large population of C. tubicula ( Towner 127), but with no evidence of hybrids. Likewise, no hybrids were apparent in a situation near the Glass Mountains in Brewster Co., Texas, where C. tubicula, C. lavandulifolius, and C. hartwegii subsp. pubescens were all discovered within a 0.3 mile stretch of graded roadside. To the north of Roswell, New Mexico, C. tubicula was observed growing within a few miles of C. hartwegii subsp. filifolius in apparently identical habitats. It can be inferred that these two taxa occasionally come into contact. Their similarity in pubescence might well be a result of some past genetic exchange, although I have seen no signs of current hybridization. 4b. Calylophus tubicula subsp. strigulosus Towner, subsp. nov. Differt a subsp. tubicula ovario et caulibus superis minute strigulosis, foliis linearibus vel anguste lanceolatis, et floribus plerumque rubescentibus vel purpurascentibus. Minutely grayish-strigulose on the ovary and upper stems, sometimes through- out, glandular pubescence generally absent. Leaves linear to narrowly lanceolate, 10-35 mm long, 0.8-3 mm wide, often shallowly serrulate. Flowers commonly fading reddish to purple. Self-incompatible. Gametic chromosome number, n=7 TYPE: MEXICO. NUEVO LEON: Along Highway 60, 2 mi W of Galeana Jct., dry rocky open range, 1,700 m, 5 July 1963, McGregor, Harms, Robinson, del Rosario, & Segal 119 ( DS-504949, holotype; KANU, SMU, isotypes). Distribution (Fig. 17): Uncommon in rocky open sites and canyons in rela- tively dry montane areas, sometimes in pine forest; southernmost Coahuila, south- central Nuevo León, and southeastern Tamaulipas. From ca. 1,500 to 2,300 m elevation. Records of flowering include July and August. Representative specime ns examined: :XICO, NUEVO LEÓN: 15 mi SW of Galeana, Mueller & Mueller 464 (F, TEX). Hacienda 1977] TOWNER—CALYLOPHUS 99 Pablillo, Galeana, Taylor 68 (ARIZ, DS, MO, NY, TEX, UC). On canyon wall, 5,400 ft, municipality of Galeana, Chase 7745 (ARIZ, F, NS NY). Ca. 35 mi S of Ca leana towards Ascención, Straw & > Forman T ( RSA ). pm recepi s Nuevo León Highway 60 W of Galeana, Sanderson 287, 288 in part (TEX). COAHUILA: SE of Saltillo, Clark 6710 (MO). Fraile, 59 km S of Saltillo, Stanford et al. 242 (DS, MO, NY, UC). rAMaULIPAs: 3 mi N of Miquihuana, Stanford et al. 2472 (DS, NY, RSA, WTU ). Jaumave Valley, Nelson 4461 (US). The ecological distribution of Calylophus tubicula subsp. strigulosus, which occurs in montane areas of northeastern Mexico, often with pines, contrasts sharply with that of subsp. tubicula. The feature in common between the two distributions is perhaps aridity, although the Mexican sites would appear to be more mesic. Several collections assigned to this subspecies show similarity to subsp. tubicula. Typically such resemblance consists of combinations of strigulose and glandular pubesence or of glandular pubescence and leaves as found in subsp. strigulosus. Of the collections cited above, intermediacy is shown by Clark 6710, Nelson 4461, and Stanford et al. 2472. Populations in the area of the type lo- cality seem the most distinct from subsp. tubicula. Two plants cultivated from seed had 3 bivalents and 2 rings of 4 chromo- somes at meiosis, each thus being heterozygous for 2 translocations. These speci- mens showed morning anthesis similar to that seen in subsp. tubicula. No field collections or observations were made. Introgression between C. hartwegii subsp. hartwegii and C. tubicula in north- ern Mexico may have given C. tubicula subsp. strigulosus its distinguishing characteristics, Such hybrid origin may have occurred quite recently since the two forms still occur in close proximity and in similar habitats in the Sierra Madre Oriental. The strigulose pubescence, greater prominence of anthocyanins, and narrow leaves of subsp. strigulosus in comparison to subsp. tubicula all represent points of similarity to the local populations of C. hartwegii subsp. hartwegii. Of the few collections we have from this area, one set seems to be fully inter- mediate with C. hartwegii, having a floral tube length of 19-21 mm [15 mi SW of Galeana, Nuevo León, Mueller & Mueller 464 (F, TEX) ]. Section II. Calylophus. "—— 4 Hist. Nat. Vég. Phan. 4: 349, 1835. Oenothera subgen. Calylophus (Spach) 1840. Merioli iat ex SE D uo n 1840; Raf., Amer. Monthly Mag. & Crit. Rev. 4 819, nom. nud. Raf., J. Phys. Chim. Hist. Nat. Arts 89: 259. 1819, nom. nüd. Herbaceous to suffrutescent perennials or annuals, 1-8 dm high, glabrous to strigulose or strigulose-canescent. Leaves 1-9 cm long, subentire to spinuose- serrate. Inflorescence dense, with buds usually crowded at the stem apex; buds squarish in cross-section. Flowers opening near sunrise. Floral tube funnelform and somewhat squarish in cross-section distally, tubular in proximal one-third to one-half of length, 2-20 mm long. Sepals with keeled midribs. Petals suborbicular to obcordate. Stamens biseriate, the episepalous filaments about twice as long as the epipetalous filaments. Capsule often tardily dehiscent, sometimes slightly recurved. 100 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TYPE SPECIES: Calylophus serrulatus ( Nutt.) Raven. 5. Calylophus berlandieri Spach, Ann. Sci. Nat. Bot., sér. 2, 4: 272. 1835. Herbaceous to suffrutescent perennial or annual arising from a woody caudex; stems one to many, simple to moderately branched, subdecumbent to erect, 1-8 dm high, glabrous to strigulose or strigulose-canescent, especially above. Leaves sessile or indistinctly petiolate, sometimes early deciduous below, spreading to more or less ascending, linear to narrowly lanceolate or oblanceolate, often folded lengthwise, 1-9 cm long, 0.1-0.9 cm wide, usually not much reduced up the stem, the abaxial surface glabrous to strigulose-canescent, especially at the base, the adaxial surface glabrous to sparsely strigulose, the tip acute, the base attenuate, the margin subentire to spinuose-serrate; fascicles of small leaves to 20 mm long often present in nonfloriferous axils; lowest stem leaves narrowly oblanceolate to oblanceolate or even spatulate. Inflorescence more or less compact, with buds usually crowded at the stem apex and one to several flowers fresh at one time, sparsely and minutely strigulose to densely strigulose-canescent; buds squarish in cross-section. Floral tube funnelform and somewhat squarish in cross-section distally, tubular in proximal one-third to one-half of length, 5-20 mm long, 3-14 mm wide at the throat in pressed specimens, subglabrous to strigulose-canescent without, especially along the midribs, within glabrous distally and minutely pubescent to strigulose basally, pale yellow green, sometimes blue black within, rarely fading pinkish. Sepals 4-12 mm long, 2-7 mm wide, with subulate free tips 0-4 mm long, with raised or keeled midribs, subglabrous to strigulose-canes- cent, pale yellow green, occasionally with red midribs and tips, only rarely fading pinkish. Petals suborbicular to obovate-truncate or obcordate, 6-25 mm long, 7-30 mm wide, occasionally becoming orangish to purplish on wilting, highly ultraviolet-reflective, but with large basal ultraviolet-absorptive spot. Stamens biseriate; episepalous filaments 2-8 mm long, the epipetalous filaments 1-4 mm long; anthers 2-7 mm long; pollen fertility normally 85-100%. Style 9-30 mm long, glabrous above and glabrous to minutely pubescent basally; stigma discoid to squarish, 1-3 mm broad, sometimes blue black, generally exserted to the ends of the anthers or beyond; ovary 5-20(-27) mm long, 0.5-1.5 mm wide, minutely strigulose to strigulose-canescent. Capsule 10-35 mm long, 1-2 mm wide, hard and thick walled, completely and often tardily dehiscent, sometimes slightly re- curved; seeds 1-1.8 mm long, sharply angled, truncate at the apex. Self-incompat- ible. Gametic chromosome number, n — 7 TYPE: UNITED STATES. TEXAS: On shore of Espiritu Santo Bay, probably in present Calhoun Co., March or May 1829, Jean Louis Berlandier 539 — 1919 ( P, holotype; GH, PH, isotypes). This locality is given on the Gray Herbarium sheet and by Spach (1835b: 338). The date was determined from information given by McKelvey (1955) Distribution (Fig. 18): Open, moderately dry areas on a variety of well- drained soils, frequently calcareous, in southeastern Colorado, southwestern Kansas, western and central Oklahoma, eastern New Mexico, Texas (except in the northeast), Louisiana, north-central Coahuila, northern Nuevo León, and 101 TOWNER-—CALYLOPHUS 1977] | | | | | TT " | * | I | 7 P, ËA j f ; (f) f eds]. | ] | { | | PER 1 ` I \ s | ^ f | I | ig { Distributions of Calylophus berlandieri subsp. berlandieri ( dots), C. berlandieri 8). Ficure 18. subsp. pinifolius (squares), and intergrades between the two subspecies (open circle 102 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 perhaps northern Tamaulipas. From sea level to ca. 1,200(-1,800) m elevation. Flowering March to September. An outcrossing species, Calylophus berlandieri is characterized by self-in- compatibility, large flowers, and the absence of permanent complex structural hybridity. Incompatibility systems of the S-allele type were demonstrated by Linder & Brun (1957) in plants of this species. I tested 24 plants from 6 popu- lations and found all to be self-sterile. The yellow flowers are morning opening, open throated, and have petals with conspicuous contrast patterns in the long- wave ultraviolet region (Fig. 14). A variety of diurnal and matinal insects, in- cluding skippers, small butterflies, bees of various families, and beetles, were observed as flower visitors. These groups probably make varying contributions to pollination at different localities, but no highly specific relationship appears to have evolved with any particular type of insect. Analysis of meiotic pairing in this species demonstrated a high frequency of translocation heterozygosity. Of 68 plants analyzed from 45 populations, 56 or fully 82% were heterozygous for 1 to 5 reciprocal translocations. The most fre- quent classes of chromosomal types were those with 1 or 2 interchanges, but several plants had as many as 5 translocation heterozygosities. No permanent structural hybridity was proven in this self-incompatible species, but the high frequency of heterozygotes implies that some developmental or selective effect probably reduced the numbers of homozygotes. As reported by Towner & Raven (1970), Berlandier’s type possesses highly fertile pollen, indicative of the pair-forming, outcrossing species in this group. A later examination of the pollen from isotypes of C. drummondianus revealed that those plants were only half-fertile. Further study of the specimens and their localities made it obvious that they were complex structural heterozygotes, thus belonging with C. serrulatus. This leaves C. berlandieri as the earliest available name for the outcrossing species. The taxonomic separation of C. berlandieri from C. serrulatus on the basis of differences in cytology and breeding systems rationalized a formerly confusing situation in regard to geographical variation. As previously interpreted, these forms presented a complex pattern of countering trends in variation of floral and vegetative parts. In my treatment, there is less conflict in variation patterns, al- though the situation is still not simple. The two species exhibit parallel geo- graphic variation, and both display a wide range of statures, leaf dimensions, and flower sizes. The parallelisms are seen primarily in vegetative characters, whereas the floral characters generally differ between the two species. Calylophus berlandieri is polytypic, with two well-differentiated morphologi- cal races. Calylophus berlandieri subsp. pinifolius, a central Texas subspecies, intergrades with C. berlandieri subsp. berlandieri in southern and west-central Texas and, to a lesser extent, in the boundary between the Edwards Plateau and the coastal plain. Populations of this species occasionally occur together with C. tubicula, C. hartwegii, and C. lavandulifolius, but I have seen no evidence of interbreeding. Numerous test crosses performed on those combinations only rarely produced 1977] TOWNER—CALYLOPHUS 103 viable offspring, and these were completely pollen-sterile. Sympatric occur- rences with C. serrulatus are infrequent, and will be discussed in the section on that species. KEY TO SUBSPECIES a. Stems several to many, subdecumbent to ascending, 1—4 dm tall; leaves 1-4 cm long E OP a. subsp. berlandieri aa. Ste ms one te ‘several, suberect to erect, 3-8 dm tall; leaves. 2. 5-9 c cm long . 5b. subsp. pinifolius Ha. Calylophus berlandieri Spach subsp. berlandieri. Oenothera berlandieri (Spach) Steud., Nom. Bot, ed. 2. 2: 206. 1841. Meriolix berlandieri Spach) Walp., Repert. Bot. Syst. 2: 79. 1843. Calylophus drummondianus Spach subsp. berlandieri (Spach) Towner & Raven, M: — 20: 243. 1970 Oenothera serrulata Nutt. var. typica sensu Munz, Amer. J. Bot. 16 712. 1929, 5 parte. Calylophus semiotic (Nutt.) Raven subsp. pads sd sensu Shinners, Sida 1: 338 pro parte. Oenothera serrulata subsp. serrulata sensu Munz, N. Amer. Fl., ser. 2, 5: 141 1965, pro des. Oenothera serrulata Nutt. var. pinifolia Engelm. ex A. Gray sensu Munz, Amer. J. Bot. 16: 715. pro parte. O. ucro subsp. pinifolia * ex A. Gray) Munz sensu Munz, Amer. Fl., ser. 2, 5: 1965, pro 5 Oconothera serrulata Nutt. var. poe ` Torr. & j Gray sensu Munz, Amer. J. Bot. 16: 714. pro parte. O. serrulata subsp. A (Torr. & A. Gray) Munz sensu Munz, N. Amer. Fl., ser. 2, 5: 142. 1965, pro parte. Perennial; stems several to many, moderately branched, subdecumbent to ascending, 1-4 dm high. Leaves more or less crowded, linear to narrowly lanceo- late or oblanceolate, 1-4 cm long, 0.1-0.6 cm wide, the margin subentire to ser- rate, and occasionally somewhat undulate; lowest stem leaves frequently oblanceo- late to spatulate. Sepals often with only slightly raised midribs, the free tips 0-2 mm long. Inside of floral tube and stigma yellowish, never black. Self-incom- patible. Gametic chromosome number, n = 7. Distribution (Fig. 18): Common on grassy prairies, plains, or low hills on sandy, gravelly, and limestone soils in relatively dry areas, frequently with Pro- sopis, Quercus havardii, and Opuntia, from western Las Animas Co., Colorado, Seward, Meade, and possibly Reno cos., Kansas, south through eastern New Mexico, the Texas Panhandle, and western Oklahoma to Culberson, Ward, and rane cos., Texas, thence southeast near the Pecos and Rio Grande rivers to the Gulf Coast, becoming widespread on the Coastal Plain north to Milam Co., Texas; also occurring in the Santa Rosa Mts. of northern Coahuila and in north- ern Nuevo León. From sea level near the Texas coast to ca. 1,200 m ( Rita Blanca Lake, Hartley Co., Texas), with one record at ca. 1,800 m elevation (12 mi S of Trinidad, Las Animas Co., Colorado). Flowers March to September. Representative specimens examined: UNITED SrArEs. COLORADO: Las Animas Co.: 12 mi S of Trinidad, Brenckle 48184 1 KANSAS: Meade Co.: Just S of spillway at Lake Larrabee, Meade Co. State Park, KANU). Reno Co.: Hutchinson, Smyth 40 (US). Seward Co.: 14 mi NE of Liberal, 3 11206 1 NEW MEXICO: De Baca Co.: 35.5 mi S of Ft. Sumner, . 2 130 (DS). Eddy Co.: 6 mi SW of White's City, Munz & Gregory 23359 (UC, RSA). 11.4 mi SW of White’s City, Towner 21 (DS). Eddy or Lea Co.: Shinneries E of Carlsbad, Goadding 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 4586 (ARIZ). Lea Co.: S of Jal, 5 0 14482 (DS). Quay Co.: Porter, Suggs in 1942 (NMC). Roosevelt or Lea Co.: Between Tatum and Portales, Goodding i in 1937 (ARIZ). OKLAHOMA: Beaver pa a8 mi m of Beaver City, Stevens 366 ( DS, NY, OKL, OKLA, SMU, US). Custer Co.: 11 and 1 W of Weatherford, Waterfall 1593 (ARIZ, NY). Harmon Co.: l.l mi W of Vinson, p mima 87 (DS). Harper Co.: Supply, Demaree 12392 (GH, OKL, POM, SMU). Jackson Co.: N bank of Red R., SW of Eldorado, Towner 138 (DS). 8.8 mi W of Elmer, Towner 139 (DS). Kiowa Co.: 17.8 mi N of Altus, Towner 78 (DS). Roger Mills Co.: Antelope Hills, Goodman 2618 (MO, NY, OKL, POM). Texas Co.: ll mi E of Hardesty, Stephens & Brooks 21758 (DS). Tillman Co.: 2.6 mi W of Tipton, Towner 140 J Aransas Co.: Aransas Bay, Berlandier 567 = 1957 (GH, MO, POM, RSA). Armstrong Co.: 15 mi S of Claude (Palo Duro Canyon), Stephens & Brooks 25469 (DS). Bailey Co.: 2 mi S of Muleshoe, Ferris & ege 3397 (DS, MO, NY). Bastrop Co.: Bastrop Park, pote 101 (TEX). Bexar Co.: 20 mi S San Antonio, Metz 678 (NY, RM). Brooks Co. 4 mi N of Hebbronville, Towner 57 (DS). 14 mi S of Falfurrias, Towner 60 (DS). 5 mi iN of Falfurrias, García 49 (OKLA, SMU, T EX). Caldwell Co.: W of Luling, Crockett 218 (LL). Calhoun Co.: Port O'Connor, Tharp in 1930 (TEX). 7.5 mi W and 5.6 mi N of Port O'Connor, Towner 179 (DS ). 2.4 mi E of Seadrift, Towner 180 (DS). Callahan Co.: Ca. 17 mi SE of Abilene, Henderson 64-52 (DS). Childre sss Co.: Estelline, Reverchon 4307 (GH, MO, NY, POM, US). Concho Co.: 2.5 mi W of Eden, Munz & Gregory 23426 (RSA). Crockett Co.: 25 mi W of Ozona, McVaugh 8209 (LL, TEX). Dickens Co.: On side of canyon, S of U.S. 82, Lundell 12979 (LL, TEX, US). Donley Co.: Jericho, Demaree 12439 DS, NY, OKL, PH, POM, TEX). Duval Co.: 15 mi E of Hebbronville, Sandoval & McCart 7982 (OKL A, TEX). Gaines Co.: 3 mi S of Seagraves, Tharp in 1941 p H, SMU, TEX). Garza Co.: 2.5 mi E of Post, Raven & Gregory 19304 (DS). Glasscock Co.: 3 mi E of Garden City, Munz & Gregory 23422 (RSA, UC). Goliad Co.: Near Goliad, Williams 9 (F, PH). Harris Co.: Spring, Gentry 865 (RM). Hartley Co.: 10.4 mi N of Channing, Roberts 35 (DS). Hemphill Co.: 7 mi ENE of Canadian, Delso 122 (DS). Hidalgo Co.: 25 mi N of Edinburg, Slant 811 (NY). Hutchinson Co.: Near Stinnett, McFarland 13 (OKL, RM). Irion Co.: rnhart, Warnock T343 (TEX, US). Jackson Co.: 13.4 mi W Palacios, Towner 176 (DS E Hogg Co.: 9.7 mi E of Hebbronville, Towner 56 (DS). Jim Wells Co.: 20 mi N of Premont, Cabrera 102 (TEX). Kenedy Co.: El Toro I, Tharp 49123 (in part, ue a mixture with C. serrulatus; F, MO, OKLA, PH, POM, TEX, UC, US). 7.5 mi S of Riv Towner 186 (DS). 17.8 mi N of Raymondv ille, Towner 192 (DS). Kleberg Co. : 0.8 mi Ww of Riviera, Towner 185 (DS). Kinney Co.: 9 mi W of Brackettville, McVaugh 7685 (DS, F, SMU, TEX). La Salle Co.: Near Cotulla, Small & Wherry 11941 (NY ). Lee Co.: Giddings, Hall 209 (F, GH, MO, NY, “POM, US). Lipscomb Co.: 23.7 mi N of Canadian, DM ll 10414 (DS). Live Oak Co.: Mikeska, Owens & Parks 2407 (MO). Loving Co.: Between Mentone and Wink, Warnock in 1952 (LL, SMU). Lubbock Co.: N of Lubbock, Demaree 7528A (DS, RSA, SMU, TEX). Motley Co.: 5.2 mi WSW of TR d Shinners 18668 (OKLA, SMU). Nueces Co.: Bishop, Eifrig in 1926 (POM). Ochiltree Co.: 11.1 mi SE of Perryton, Towner 158 (DS). A Pinto Co.: 19 mi W of Mineral Wells, Warren "3 (DS). Pecos Co.: 30 mi W of Sheffield, Munz 13290 (DS, POM, UC). Potter Co.: 1 mi N of Canadian R. on Highway 287, Jefferson & Jefferson 2676 (DS, F, MO, NEB, NY, RM, SMU, UC, US, WTU). .3 mi N and 2 2.0 mi W of Amarillo, Towner 91 (DS). Randall Co.: Palo Duro State Park, Cory 13036 (LL, SMU). Refugio Co.: 8.9 mi W E Refugio, ps 1 5 6831 (SMU). San Patricio Co.: Near Mathis, McKelvey 1726 (GH). Taylor Co.: 3 mi S of Camp Barkeley, Tolstead 7096, 41983 (MO, NEB, NY, POM, SMU, i. Terrell TAM 13 mi S of Sheffield, Webster 130 (TEX). Tom Green Co.: 7% mi S of Christoval, Cory 50569 (NY, SMU) Travis Co.: Austin, Tharp in 1938 (SMU, UC). Uvalde Co.: 6 mi SE = Uvalde, Munz 13316 (POM). Val Verde Co.: Ca, 1.9 mi from Del Rio, Traverse 2162 (SMU, TEX). Victoria Co.: Inez, Palmer 9137 (DS, US). Ward Co.: 3 mi ENE of Monahans, McVaugh 8186 (DS, GH, LL, TEX). Webb Co.: 7 mi N of Laredo, um ^y 129 (SMU, TEX). Wheeler Co.: 11.5 mi E of Shamrock, Rowell 10080 (DS, RSA). Wilbarger Co.: 20 mi N of Vernon, Towner 77 (DS). Willacy Co.: 1*4 mi from shore at Port Mansfield, Webster d» Wilbur 3074 (SMU, US). Wilson Co.: 5 mi N of Stockdale, Munz & Gregory 23443 (RSA, UC). Winkler Co.: 11 mi W of Kermit, Raven & Gregory 19228 (DS). Wise Co.: 3 mi N of 5 Whitehouse 15278 (SMU). Yoakum Co.: 4.7 mi N of Bronco, Towner 135 (DS). Zapata Co.: 5 mi SE of San Ygnacio, Flores & Flores 147 (TEX). Counties not known: From Bejar (San eer to Austin, Berlandier 479 = 1829 (GH). From Matamoros to Goliad. Berlandier 1048 = 2478 (GH, MO, PH). — — — 1977] TOWNER—CALYLOPHUS 105 ICO. COAHUILA: Santa Rosa Mts., Marsh 1354 (F, OKLA, SMU, TEX). Muüzquiz, Marsh 110 (F, OKLA, TEX). Hacienda M: ariposa, Mcpo. of Müzquiz, Wynd & Mueller 264 (ARIZ, MO, NY, US). Summit of La Cuesta Malena Mts., Reveal et al. 2594 (DS). NUEVO LEON: Lampazos, Edwards 356 (F). This relatively short-leaved and low-statured subspecies occurs over an ex- tensive range on the plains of Texas and adjacent states. It is common in the Texas Panhandle and along the Gulf Coast, and it is also found locally in sandy areas of West Texas. As I have defined it, Calylophus berlandieri subsp. ber- landieri incorporates clements from each of the three subspecies of “Oenothera serrulata” recognized by Munz (1965: 141-142). For instance, the extremely narrow-leaved plants formerly known as Oenothera serrulata subsp. pinifolia are clearly variants which can actually be found along with broader-leaved plants in populations of either subspecies of C. berlandieri. My treatment of these forms is similar to that of Shinners (1964), who did not recognize subsp. pinifolia, viewing it as merely the extreme in a wide range of variation, the latter due to “spontaneous mutation.” The narrow-leaved plants are most frequently found in areas of highly calcareous soil, including gypsum, and they occur in the more arid portions of the range of C. berlandieri. Thus their presence may well be due not to “spontaneous mutations,” but to edaphic selection factors. In the past, narrow-leaved individuals of C. berlandieri subsp. berlandieri were often assigned to Oenothera serrulata var. pinifolia. Examples are the fol- lowing: 5 mi N of Stockdale, Wilson Co., Texas, Munz & Gregory 23443 (UC, RSA). 2 mi S of Muleshoe, Bailey Co., Texas, Ferris & Duncan 3397 (DS, MO, NY). Narrow-leaved individuals are slightly more frequent in C. berlandieri subsp. pinifolius, although they are not representative of that taxon as a whole. The distribution of C. berlandieri subsp. berlandieri is divided into a Coastal Plains section and a Central Plains section. These two series of populations are connected only tenuously, this through a narrow zone along the Rio Grande southwest of the Edwards Plateau. The coastal region is less severe in climate than is the central region, but both areas are semiarid. Considering the breadth of these separate ranges, their distinctness, and their climatic differences, one might expect the two series of populations to have diverged morphologically. However, this does not appear to have been the case, as they show completely overlapping ranges of variation in all characters which I have examined. In spite of the morphological similarity of the two series, meiotic pairing in hybrids suggests that considerable cytological divergence may have taken place. Nine hybrids between progeny of Rowell 10414 (Lipscomb Co., Texas) and progeny of Bohart & Thorp 650928-1 (Victoria Co., Texas) showed I/ KI pollen stainability of approximately 40%. Meiotic determinations of 2 of the hybrids showed chains of 14 chromosomes, indicating the presence of at least 6 reciprocal translocation differences between the parents. Similar data were obtained from crosses of Rowell’s collection with Towner 185 (Kleberg Co., Texas). The frequency of translocation heterozygosity in natural populations of Caly- lophus berlandieri subsp. berlandieri was found to be extremcly high. Of 38 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 plants examined, including some grown from field-collected seed, only 6 formed 7 bivalents at meiotic metaphase. The remaining 84% showed multivalent forma- tion in various degrees, indicating structural heterozygosity. The average num- ber of translocations per plant was 2.0. Flower visitation was observed at one site in Potter Co., Texas (Towner 91). Anthesis occurred shortly before sunrise, at which time several hawkmoths ( Hyles lineata) visited flowers for nectar. At sunrise and afterwards, skippers, small butterflies, and bees of small to medium size, e.g., Evylaeus and Agapostemon, collected nectar and pollen from the flowers. Pollination was perhaps most ef- fective with the anthophorid bees ( Melissodes and Anthophora), which were few in number, however. Oligolectic halictids probably made some contribution, since they were common and because their appearance coincided closely with anthesis times. Intergradation occurs between C. berlandieri subspp. berlandieri and pini- folius in several areas. Along the eastern escarpment of the Edwards Plateau, few natural intermediates are found, apparently because the respective habitats of the two forms are separated by a relatively sharp geographical discontinuity. On the southern and western sides of the Plateau, intermediate forms are much more frequent. There the geographical changes are more gradual, and broad zones of hybridization are evident. On the western side of the range of C. ber- landieri subsp. pinifolius, numerous plants intermediate in stature and leaf length are found: 25 mi W of Ozona, Crockett Co., Texas, McVaugh 8209 (LL, TEX). 3 mi S of Camp Barkeley, Taylor Co., Texas, Tolstead 7096, 41983 (MO, NEB, NY, PO, SMU, UC). 3 mi E of Garden City, Glasscock Co., Texas, Munz & Gregory 23492 (RSA, UC). Similarly, collections intermediate between C. ber- landieri subsp. pinifolius and the Rio Grande Valley populations of subsp. ber- landieri are relatively frequent. Several examples are as follows: Ca. 1.9 mi from Del Rio, Val Verde Co., Texas, Traverse 2162 (SMU, TEX). 9 mi W of Brackett- ville, Kinney Co., Texas, McVaugh 7685 (DS, F, SMU, TEX). 6 mi SE of Uvalde, Uvalde Co., Texas, Munz 13316 ( POM ). Over much of its distribution, C. berlandieri subsp. berlandieri occurs with or near populations of species of sect. Salpingia. In most cases of sympatry a few plants of one form are found scattered in or near a colony of the other, and only rarely are both forms common at any locality. The taxa I observed growing together with C. berlandieri subsp. berlandieri were C. hartwegii subsp. filifolius in De Baca Co., New Mexico and C. hartwegii subsp. pubescens in the Texas Panhandle. Also occurring with C. berlandieri subsp. berlandieri in the Pan- handle and in western Oklahoma, but more distinct ecologically, are C. hartwegii subsp. fendleri and C. lavandulifolius. In West Texas there is some likelihood of contact between C. tubicula and C. berlandieri subsp. berlandieri, although their edaphic restrictions seem to reduce this possibility severely. Lastly, local sympatry may also occur with C. hartwegii subsp. maccartii in the lower Rio Grande Valley, since the edaphic and geographical ranges of the two forms over- lap there. Me] ~J -1 TOWNER-—CALYLOPHUS 107 5b. Calylophus berlandieri Spach subsp. pinifolius (Engelm. ex A. Gray) Towner, comb. nov.—Fic. 5. VENE "s Nutt. var. d Engelm. ex A. Gray, Boston J. Nat. Hist. 6: 189. 1850; , Am ]. Bot. 16: 1929. Meriolix serrulata (Nutt.) Raf. (var.) pinifolia (E nge Na ex v Gray ) di F Torrey Bot. Ch x 23: 187. 1896. Oenothera serrulata subsp. pinifolia ( Engelm. ex A. Gray) Munz, N. Amer. Fl., ser. 2, 5: 141. 1965. Oenothera captlifoltà Scheele, Linnaea 21: 576. 1848. "Meriolix capillifolia (Scheele) Small, S. E. 846, 1335. 1903. type: United States, Texas, Comal Co., New Braunfels, ril (18462). Ferdinand Roemer (not located ). baa jo Small, F]. U.S. 846, 1335. 1903. TYPE: United States, Texas, Edwards o Water Pu - June 1895, R. T. Hill ( NY ut she Sa Rydb. ex Small, Fl. S.E. U.S. 846. 1335. 1903. Oenothera ser- rulata Nutt. var. ace H. Lév., Monogr. Onoth. 336, 339. ee (lectotype: Hel- ler 1600, MO). United States, Texas, Kerr Co., Kerrville, 19-25 April 1894, A. A. Heller 1600 (Ny. holotype; ARIZ, MO. NEB, NY, PH, POM, RM, SMU, UC, otypes ) US, iso Oenothera serrulata Nutt. var. drummondii Torr. & A. Gray sensu Ma Amer. J. Bot. 16: 714. 29, pro parte. O. serrulata subsp. drummondii (Torr. & A. Gray) Munz sensu Munz, N. Amer, Fl., ser. 2, 5: 142. 1965, pro parte. um serrulata Nutt. var. drummondii 'Torr. & A. Gray : qu Munz sensu Munz, Amer. 16: 714. 1929, for the most part, excluding the ty Calls serrulatus ( Nutt. ) Raven var. spinulosus (Torr. & A. ‘Gan Shinners sensu Shinners, 964, pro parte. Calylophun drummondianus Spach subsp. drummondianus sensu Towner in Correll & Johnston, Man. Va „Pl. Texas 1123. 1970, all, except for the typ — Annual to short-lived perennial; stems one to several, simple or sparsely branched, suberect to erect, 3-8 dm high. Leaves well spaced, linear to narrowly oblanceolate or narrowly lanceolate, 2.5-9 cm long, 0.2-0.9 em wide, the mar- gin remotely serrulate to spinuose-serrate; lowest stem leaves narrowly ob- anceolate. Sepals with conspicuously keeled midribs, with free tips 0.5-4 mm long. Stigma and inside of floral tube frequently deep blue black in certain populations. Self-incompatible. Gametic chromosome number, n = 7 — TYPE: UNITED STATES, TEXAS: Comal Co., rocky prairies, New Braunfels, April 1846, Ferdinand Lindheimer 37 = 394 (GH, holotype; DS, MO, NY, PH, RSA, US, isotypes ). Distribution (Fig. 18): Common on prairies and in open places in oak savan- na, on rocky, clay, or sandy soils, often calcareous, from Blaine and Lincoln cos., Oklahoma, south through a narrow portion of north-central Texas to central Texas, where it is widely distributed, especially on the Edwards Plateau; also occurs locally in western and southern Louisiana. Elevational distribution from near sea level (Sulfur, Calcasieu Parish, Louisiana) to ca. 900 m (Sonora, Sut- ton Co., Texas). Flowers mostly from March to June. 5 specimens examined: Unr 5 MISSOURI: Jackson Co.: Sheffield (introduced), Bush 328 (F. GH). OKLAHOMA: Blaine Co.: 3 mi W of G 8 Id, Hop vis = van Valkenburgh 4131 (OKL). Caddo Co.: Rim ‘of Devil's Cub Hopkins et al. 309 (DS, F, MO, OKL, OKLA, RM, SMU, UC, WTU). Canadian Co.: 2 mi W of El Reno, Munz 4 rap penn 23505 (RSA, UC, WTU). Cleveland Co.: Norman, Demaree 12473 (MO, NY, OKL. POM, RM, SMU). Cotton Co.: 6.9 ee E of 2 1 141 (DS). 9 mi W of Comanche, Towner 142 (DS). Custer Co.: za. 1.5 mi W of Custer, Towner 154 (DS). Lincoln Co.: 3.2 mi S of Perkins, Towner 149B 1 Logan Gas. 15 mi W of Guthrie, Raven & Gregory 19462 (DS). 5.4 mi W of Guthrie, Towner 151 (DS). McClain Co.: Blanchard (Johnson's Pasture), Demaree 13094 (MO, NY, 10S ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 OKL, PH, POM, TEX, UC, US). Oklahoma Co.: 6 mi W and 6 N of ar 758 City, W aterfall 1423 Bee. POM). Seminole Co.: SE of Konawa, Robbins 2458 (O UC). = ^ C) A ° *phens : of Comanche, Waterfall 3679 (NY, OKL). Texas: Austin Co.: £ e Fisher 3846 iF). Bastrop Co.: S of Bastrop, Lundell & Lundell 10355 (ARIZ, LL, POM, >). Bell Co.: 5 mi S of Temple, Wolff 4037 (F). Be xar Co.: San Antonio, Clemens & Clemens 686 (NY, PH, POM, RM). Blanco Co.: 2 mi S of Blanco, Tharp et al. 17T195 (RM, SMU). 5.5 mi N of Johnson City, Towner 68 (DS). 32.7 mi NW of San Marcos, Towner 66 (D S). Brown Co.: 8.2 mi S of Brownwood, Towner 72 (DS). Brownwood, Ewing 26 (LL, SMU, TEX). Burnet Co.: 1 Fisher 40065 (ARIZ, dei TEX). Coke Co.: 2 mi E of e Lee, Shinners 31773 (SMU). Coleman Co.: 14.3 mi N of Coleman, Towner 74 (DS). Comal Co.: New Braunfels, Lindheimer 809 ( ARIZ, DS, F. G = MO, NMC, NY, OKL, PH, TEX, UC, US). 20.9 mi NW of San Marcos, Towner 64 (DS). Comanche Co.: Round Top Mt., Eggert n vds ieu Coryell Co.: 13 mi E of Gatesville, Jackson 4 (LL, SMU, TEX). Crockett 0. N of Juno, Warnock d (LL, TEX). Dallas Co.: White Rock Lake, etel d ‘tae I 8531 (DS, GH, LL, POM, SMU). Dimmit Co.: Carrizo Springs, Palmer 33732 (MO). Ellis Co.: 1% mi N of Midlothian, Cory 53327 (DS, KANU, RM, SMU, UC). Erath Co.: 3.5 mi W of Stephensville, Gould 5644 (RSA, SMU, UC). Fayette Co.: La Grange, Hanisch in 1935 (TEX). Frio Co.: 6 mi NE of Pearsall, Lundell 13628 (LL, TEX). Gillespie Co.: Bear Mt., Correll & Correll 12763 (LL, SMU). Hamilton Co.: Hamilton, Tharp in 1941 (GH). Harris Co.: Houston, Fisher 3481 (F). Hays Co.: 5 W of San Marcos, Gregory 419 (DS, RSA, US). 5.1 mi NW of San Marcos, Towner 62 )$). 7.5 mi NW of San Marcos, Towner 63 (DS). Hill C5. 8.5 mi NE of Hillsboro, Shinners 12494 A 1 Co.: Grandbury, “Naples 8 7174 (US). Irion Co.: 30 mi N of Barnhart, Rat & Gregory 19210 (DS). Karnes Co.: 3 mi SE of Karnes City, Johnson 857 (RSA, TEX). Kendall Co.: 19 mi S of Fre dede Munz & Gregory 23438 VTU). Kerr Co.: 4 mi SW of Kerrville, Cory 51 1 (Ds, SMU). Kimble Co.: No locality, Tharp 43-738 (TEX, uc). ae Co.: Ca. 30 mi SE of Brackettville, Strother 240 (DS, SMU). Lampasas Co.: S of Lampasas, W "hitehouse 15384 (SMU). Llano Co.: No locality: Lundell & Landen 9050 (DS, GH, LL, POM, SMU). 1.7 mi N of Llano, Towner 69 (DS). Medina Co.: Hondo, Pilsbry in 1903 (PH). Menard Co.: 10.3 mi N of Menard, Raven & Gregory 19273 (DS). McClennan Co.: Betwee en Waco and McGregor, York 46074 (TEX, UC). McCulloch Co.: 9 mi SE Brady, Munz & Gregory 23431 WTU). McMullen Co.: No locality, Schultz 64 (US). Mills Co.: ^ Goldthw. aite, Ferguson z 9] ( MO, PH, TEX, UC). San Saba rs 4 mi W of Pontotoc, Jones 24 (LL, 9 Sutton Co.: Sonora, Tharp in 1931 (TEX). Tarrant Co.: Lake Como, Ruth 30 (F). Travis Co.: 9 mi W of Oak Hill, Lundell & 2 95 8898 (DS, GH, LL, NY, POM, RM, aes UC). Uvalde Co.: 20 mi N of Uvalde, Graves 9 (RM, RSA). Val Verde Co.: Devil’s R. N of Del Rio, te in 1903 (PH). Washington Co.: No locality, Brackett in 1938 (GH). Williamson : » locality, York 46193 (TEX). Louisiana: Acadia Parish: Prairies near Crowle N e C 11741 (NY). Calcasieu Parish: m Palmer 7719 (MO). St. Mary Parish: Near Berwick, Small & Wherry in 1925 (NY). — = = 5 — C P As with Calylophus berlandieri subsp. berlandieri, subsp. pinifolius in- corporates portions of several of the taxa recognized by earlier authors. It cor- responds rather closely with Oenothera serrulata subsp. drummondii except for the inclusion of narrow-leaved individuals and exclusion of complex heterozygotes. As mentioned earlier, the types seen of C. drummondianus were erroneously as- signed to the outcrossing species in a preliminary report (Towner & Raven, 1970). The types actually belong with the complex structural heterozygote C. serrulatus, and the name is reduced to synonymy. The collection used as the basis for Oenothera serrulata var. drummondii f. flava was also found to belong with C. serrulatus, having the small flowers and half-sterile pollen indicative of com- plex structural heterozygosity. Calylophus berlandieri subsp. pinifolius is distributed primarily from central Texas to central Oklahoma, inhabiting more mesic areas than does subsp. ber- landieri. Most typically it is found in calcareous, rocky soil in oak savanna. Colo- 1977] TOWNER—CALYLOPHUS 109 nies occur in open or disturbed areas in that habitat and in prairies. This sub- species is the only member of sect. Calylophus occurring on the Edwards Plateau in Texas. Earlier reference was made to the narrow-leaved variants occurring in the two subspecies of C. berlandieri. The presence of these forms in both taxa pre- vents the use of leaf proportion as a diagnostic or key character. However, C. berlandieri subsp. pinifolius is distinctly longer leaved than subsp. berlandieri, and the use of this character together with differences in stature permits an easy diagnosis of most individuals. Subspecies pinifolius generally has a slender tap- root and is difficult to maintain for more than a year in the greenhouse, leading to the inference that it is probably a short-lived perennial or annual in the field. This contrasts with subsp. berlandieri, which is distinctly perennial over most of its range. Some representatives of this subspecies possess purplish black stigmas and/or inner surface of the floral tubes. This extremely interesting character is present as a polvmorphism in many populations, but is restricted to those in south- central Texas. The dark-pigmented forms are especially frequent. in Bexar, Blanco, Comal, Gillespie, Hays, Kendall, Kerr, and Travis cos. Some examples of the variants include the following collections: Type specimen of Meriolix melanoglottis Rydb. ex Small. Type specimen of Meriolix hillii Small. 19 mi S of Fredericksburg, Kendall Co., Texas, Munz & Gregory 23438 (RSA, WTU). 7.5 mi NW of San Marcos, Hays Co., Texas, Towner 63 (DS; most plants with black stigma, some with black hvpanthium). 5.5 mi N of Johnson City, Blanco Co., Texas, Towner 68 (DS; plants with black stigma only). U.S. 57 near NW city limits of San Antonio, Bexar Co., Texas, Klein 1671-1674 (DS). New Braun- fels, Comal Co., Texas, Lindheimer S09 (ARIZ, DS, F, GH, MO, NMC, NY, OKL, PH, TEN, UC, US). 4 mi SW of Kerrville, Kerr Co., Texas, Cory 51763 (DS, SMU). Plants from populations of C. berlandieri subsp. pinifolius from Oklahoma differ modally from those in Texas in having shallower leaf serrations and more strigose pubescence on the upper stems. This is especially apparent in the north- ernmost populations, e.g, those in Logan, Oklahoma, McClain, and Cleveland cos. In regard to stature, leaf dimensions, and most other characters, these plants are identical to individuals from Texas populations of subsp. pinifolius. Some of the largest-flowered members of the species occur in these Oklahoma popula- tions. Examples of plants from this area are as follows: 5.4 mi W of Guthrie, Lo- gan Co., Oklahoma, Towner 151 (DS). Norman, Cleveland Co., Oklahoma, Demaree 12767 (OKL, POM, SMU). Blanchard, McClain Co., Oklahoma, Dema- ree 13094 (MO, NY, OKL, PH, POM, TEX, UC, US As in C. berlandieri subsp. berlandieri, this subspecies has a great deal translocation heterozygosity. Of the 30 plants which have been examined, 24 or 80% had ring or chain multivalents at meiotic metaphase I. Of these plants, those having 1 or 2 heterozvgosities were the most frequent types. The average number per Sinis was 1.6. Floral behavior has been observed in Brown Co., Texas (Towner 72) and in the greenhouse at Stanford, California. Anthesis took place shortly after sun- 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 rise. Insect visitors to flowers observed by me, and by P. H. Raven and D. P. Gregory in Hays and Kyle cos, Texas (personal communication), included small butterflies, flies, skippers [Atalopedes campestris (Boisduval)], a variety of beetles, especially cantharids and Acmaeodera ( Buprestidae), numerous small bees (Agapostemon, Augochlorella, Dialictus, Evylaeus, and Halictus), and a few large and medium-sized bees (Apis, Bombus, Megachile, Xylocopa, an- thophorids). It seems likely that a broad spectrum of insects serves as pollen vectors. Skippers, medium-sized bees, and beetles, judged by their abundance, size, and contact with the anthers and stigma, may be the primary pollinating agents for most populations. The black floral tubes and stigmas, which occur together with large ultraviolet patterns, may complement those patterns in facilitating the orientation of visiting insects. This would be especially important for any species of insect which lacked the tricolor vision of bees and could not discriminate in the ultraviolet region of the spectrum. With no evidence of hybridization, C. berlandieri subsp. pinifolius occurs to- gether with C. hartwegii subsp. pubescens in much of its range in central and west-central Texas. No other member of sect. Salpingia overlaps significantly in distribution with subsp. pinifolius. Minor geographic sympatry exists in south- ern Texas with C. hartwegii subsp. maccartii and in western Texas with C. tubicula subsp. tubicula and C. hartwegii subsp. filifolius. In these cases, local sympatry is quite infrequent or absent, probably as a result of habitat segregation. 6. Calylophus serrulatus (Nutt.) Raven, Brittonia 16: 286. 1964.—F'c. 6. . serrulata Nutt., Gen. N. Amer, Pl. 1: 246. 1818. Meriolix serrulata (Nutt.) Raf., Amer. Monthly Mag. & Crit. Rev. 4: 192. 1819. Calylophus nuttallii Spach, Hist. Nat. Vég. "Phi in. 4: 350. 1835. apas qu serrulata var. nuttallii (Spach) Torr. & A. Gray, F P Amer. 1: 501. 1840. Meriolix serrulata var. nuttallii (Spach) Walp., Repert. Bot. Syst. 2: 79. 1 Oenothera serrulata var. typica Munz, Amer. J. Bot. 16: 712. 1929. Calylophus „ (Nutt.) Raven var. serrulatus; Shinners, Sida 1: 338. 1964. Oenothera serrulata subsp. serrulata; Munz, N. Amer. Fl., ser. 2, 5: 141. 1965. Oenothera leucocara Comien ex Lehm. in Hooker, Fl. Bor. Amer. 1: 210. 1833. O. serrulata ar, douglasii Torr. & A. G ws Fl. N. Amer, 1: 502. 1840. Mertolix serrulata (Nutt. ) Raf. var. douglassii (Torr. & A. Gray) Walp., Repert. Bot. Syst. 2: 79. 1843. TYPE: ogre Saskatchewan, common on ine stone rocks on Red and Assiniboine rivers, August vid Douglas (K, lectotype). Calylophus V Spach, Ann. Sci. Nat. Bot., sér. 2, 4: 272. 1835. Oenothera serrulata drummondii Torr. & A. Gray, Fl. N. Amer. i; 502. 1840. O. spachiana Steud. Non Bot., ed. 2. 2: 207. 1841. Meriolix serrulata ( Nutt.) Raf. var. ` drummondii (Torr. & A. Gra y) Walp., Repert. Bot. Syst. 2: 79. 1843. M. 5 apara) Small, Fl. S.E. U.S. 846, 1335. 1903. Oenothera serrulata subsp. drummondii (T & A. Gray) Pang N. Amer. Fl., ser. 2, 5: 142, 1965, pro parte. Calylophus drimmondian subsp. dru nd Towner in Correll & Johnston, 9 1 Pl. xas 1123. 1970. TYPE: United States, Texas, along the Rio Braz the Texas a plain, probably between 2 os Co. and the coast, 1833, Thomas Dene (P, holotype; GH, NY, isotypes, but pt Drummond III. 79 ut PH). Gen ee serrulata Nutt. var. spinulosa Torr. & A. ME Fl. N. Amer. 1: € 1840. “sedges rrulata ( Nutt.) Raf. var. spinulosa (Torr. A. Gray) Walp., Repert. Bot. Syst. 2: 843. M. spinulosa (Torr. & A. Gray) Í Heller Ca. Herb. Frankl. š Marsh. 1: 70. 1895. Calylophus serrulatus doa Raven var. spinulo: sus (Torr. & A. Gray) Shinners, Sida 1: 339. 1964. rype: United States, Oklahoma, vicinity of the Red R., probably near present-day Choctaw Co., May-June 1819, Thomas Nuttall, oid Herb. INY, lectotype: PH, NY, isolectotypes, but not “Red River, Nuttall,” (GH), or “Arkansas,” (PH)]. Some of Nuttall’s Red River and Arkansas collections, such as the CH specimen cited, are actually 1977 TOWNER—CALYLOPHUS 111 5 berlandieri. Ne type sheet has the Leavenworth specimen cited by Torrey and Gray inted next to the Red River js of Nuttall. Meriolix Ghee iae ex Small, Fl. . U.S. 846, 1335. 1903. TYPE: United 15 755 ouri, Atchison Co., Watson, P p B. F. Bush 321 (NY, holotype; MO, OKL, types Oe Bend. serrulata Nutt. var. integrifolia H. Léy., Monogr. Onoth. 337, 339. 1908. TYPE: Inited States, probably from southeastern Colorado, 1945, third n of john C. Fremont, no. 47 MO); Munz, Amer. J. Bot. 16: 713. 1929. Oenothera serrulata Nutt, var. drummondii Torr. & A. Gr ay f. flava Pu Amer. J. Bot. 16: Uus bea TYPE: United States, Texas, Walker Co., Huntsville, B. Tharp 866 ( POM- other collections cited in the protologue are C. ewe JR pinifolius). Meriolir Khen. Rydb., Brittonia 1: 93. 1931. rype: United States, Kansas, 9 9 5 z0., along road 2 mi W of pu 8 July 1929, P. A. Rydberg & R. Imler 737 holot type; KANU, NEB, NY, Calylophus n (Nutt. ) ie var. arizonicus Shinners, Sida 1: 338. 1964. Tyee: United States, Arizona, Nav ajo Co., dry sandy riverbank 4 mi 1 ro White River. 25 June 1951, $. J. Preece, Jr. & B. 1. Turner 2692 (SMU). Calylophus australis Towner & Raven, Madroño 20: 243. 1970. rype: United States, Texas, Cameron Co., Texas route 4, 2.8 mi W of end of road at Boca Chica, 29 May 1969, Towner 187 ( DS- 612434, holotype; RSA, TEX, US, isotypes). Similar to Calylophus berlandieri. Herbaceous to suffrutescent perennial from a woody caudex; stems few to many, 1-6 dm high. Floral tube 2-12(-16) mm long, 3-12 mm wide, never blue black within. Sepals 1.5-9 mm long, 2-6 mm wide. Petals 5-12(-20) mm long, 5-15(-20) mm wide. Episepalous fila- ments 1-5(—7) mm long, the epipetalous filaments 0.5-3 mm long; anthers 1.5-4 (-7) mm long; pollen grains 30-80% aborted. Style 2-15(-20) mm long; stigma 1-2 mm broad, not exserted beyond the anthers, and often in contact with the anthers at the apex of the floral tube, never blue black; ovary 4-13 mm long. Self-compatible and highly autogamous. Gametic chromosome number, n = 7, with a ring of 14 chromosomes or a ring of 12 plus a bivalent at meiotic meta- phase I. TYPE: UNITED States. Plains along the Missouri River, probably just north of the Platte River in eastern Nebraska, April or May 1811, Thomas Nuttall (PH). From Bradbury account of the expedition (McKelvey, 1955: 115-118) and because of the immature appearance of the type, these dates seem more likely than June, the date of flowering given in the original description. Distribution ( Fig. 19): Common on plains, in grassy open areas in woods, or, rarely, in mountains, usually on sandy or rocky soils, from southern Alberta, southern Saskatchewan, and southern Manitoba to eastern New Mexico, the Texas Panhandle, and the Gulf Coast of Texas, including eastern Montana, east- ern Wyoming, eastern Colorado, North Dakota, South Dakota, Nebraska, Kan- sas, western and central Oklahoma, western and southern Minnesota, Iowa, north- western Missouri, and with outlying populations in southeastern Wisconsin, northwestern peninsular Michigan, east-central Arizona, and west-central Chihua- hua, Mexico. Elevational distribution from sea level along the Texas coast to 2,100 m (18 mi N of Rubio, Chihuahua). Flowers March to August. Repre sentative specimens examined: NADA. ALBERTA: Ne: eigan, Moss 871 (DAO, NY). saskarcuEWAN: Katepwa, JA Russell 54519 (DAO). pilot Butte, Hart in 1939 (DAO, UC). 13 mi W of —- Shumovich 38 (CAN, DAO, RM). Moose Jaw, Turner 48 (GH, NY, POM, RSA). MANITOB ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 | ç “as, YE ENT ° 22 e 2 „„ : | 3 f | [ — 2227 S. 2 os > [ N 00. WM AEXRES 01. | e | ee | °. ° / Ec "P | ete rR ° e or eere? ~~] I | Ficure 19. Distribution of Calylophus serrulatus. Brandon, Fowler in 1887 (DAO, MO, NY, US). Horton, Love d» Love 6081 (DAO, GH). Melita, Scoggan 9794 (CAN, GH). Mouth of the Qu'Appelle R., Macoun & Herriot 72,377 (F, GH, NY). UNITED STATES. MONTANA: Billings Co.: 1.5 mi S of Medora, Stephens & Brooks 13486 1977] TOWNER—CALYLOPHUS 113 (KANU). Powder River Co.: 15 mi NW of Boradus, Booth 2517 (KANU, WTU). Prairie Co.: 32 mi NW of Terry, Stephens & Brooks 23691 (DS). Sheridan Co.: Westby, Larsen 213 (GH, NY). Wheatland Co.: 12 mi S of Harlowton, Hitchcock 2429 (MO, POM, RSA). NORTH DAKOTA; Barnes Co.: Valley City, Stevens in 1934 (F, RM, UC). Divide Co.: Alkabo, Larsen 88 (CH, MO, PH). Dunn Co.: 10 mi NW of Killdeer, Stephens & Brooks 12734 (DS, KANU). Golden Valley C 0.: lmi E and 1 S of Sentinel Butte, Stephens & Brooks 23428 (DS). Slope Co.: 7 mi S of Amidon, Cutler 2615 (DS, MO, NY). Ward Co.: 9 mi NW of Minot, Stephens & Brooks 12897 (DS, KANU). SOUTH pakora: Custer Co.: 14 mi S of Pringle, Stephens & Brooks 13798 (DS, KANU). Harding Co.: 6 mi N, H W, and 2 N of Stephens & Brooks 13657 (DS, KANU). Lawrence Co.: 0.8 mi N of Spearfish, Mosquin & Mulligan 5160 (DS). Meade Co.: Near Fort Meade, Forwood p ( CAN, US). Pennington Co.: Rapid City, Rydberg 708 (F, GH, NEB, NY, US ). P Perkins Co.: 10 mi E and 1 S of Bison, Stephens 7999 (RANU, SMU). MINNESOTA! Chippewa Co.: Montevideo, Moyer in 1908 (NY, UC). Dakota Co.: 5.5 mi W of Hastings, Moore 15797 (DAO, GE Faribault Co.: Elmore, Pammel 596 (G H, MO, RM, US). Nicollet Co.: No locality, Aiton in 1891 (F, NY, POM, US). Ottertail Co.: Perham, Chandonnet in 1910, 1911 (GH, RM). Yellow Medicine Co.: S side of Granite Falls, Moore 13071 (SMU, UC). wisconsin: Pepin Co. (?): Lake Pepin, Hale 1861 (F, MO). wvowixc: Converse Co.: 15 mi N of Douglas, Stephens & Brooks 23965 (DS). Crook Co.: 5 mi NE of Hulett, Porter d» Porter 9564 (CAN, DS, RM, RSA, Uo. piu Goshen Co.: 12 mi W of Lagrange, Stephens & Brooks 22947 DS). Niobrara Co.: 10 mi N of Lusk, Mosquin d» gei 5142 (DS). Platte Co.: Guernsey, Nelson 8268 (DS, F, GH. MO, NEB, NY, POM, RM, RSA, UC, US). 3 mi W of “waw ~ = Qa — = c = z Porter 4903 (COLO, DS, GH, MO, PH, RM, RSA, SMU, TEX, WTU). NEBRAsKA: Adams 20 mi W of Hastings Mathias 308 (MO, POM). Chase Co.: 6 mi N of Imperial, Siepie ns va Brooks 11490 (KANU). Cherry Co.: Vicinity of Hackberry Lake, Dworak in 1912 (NEB). Dawes Co.: Chadron State Park, Porter & Porter 8807 (DS, RM, WTU). Garden ( Go. i S of Lewellen, Stephens & Brooks 11557 (KANU). Greeley Co.: 9 mi N of Greeley, McGregor 19344 (KANU). Hayes Co.: 14 mi W of Hayes Center, Stephens & Brooks 13958 (DS, KANU). Holt Co.: 12.5 mi S of Atkinson, Ste phens 15579 (KANU). Kearney Co.: Minden, Hapeman in 1930 (OKLA, TEX). Lili Co.: North Platte, Jones in 1925 (DS, POM). Morrill Co.: 27 mi N of Broadwater, Stephens & Brooks 13888 (DS, KANU). Saunders Co.: 4 mi S of Valparaiso, Croat 2126, 2127 (KANU, MO). Sheridan Co.: 13.5 mi S of Hay Springs, Stephens & Brooks 13839 (DS, KANU). Webster Co.: S of Blue Hill, Tolstead 11245 (NEB, UC). 10wa: Emmet Co.: No locality, Cratty, s.n. dos F, NY, PO, UC Fremont Co.: Hamburg, Bush 10313 (GH, MO, PH, POM). Lyon Co.: Gitchie Manitou State Park, 5 14234 ers UC). Palo Alto Co.: Highland Township, Hayden 10081 (PH, UC, US). Wayne Co. orydon City Reservoir, van Bruggen 2716 (UC). COLORADO: Baca Co.: 23 mi S of Walsh, Stephens & > Brooks 21788 (DS). Cheyenne Co.: 17 mi N of Kit Carson, 5 & Brooks 22662 (DS). Douglas Co.: Wolhurst, Clokey 3827 (CAN, DS, F, G MO, NY, PH, POM, RM, SMU, UC, US, WTU). "eb Co.: 4 mi SW of Limon, Ownbey 1301 (COLO, GH, ba 8 RM, UC). Kiowa Co.: mi E of Eads, Sis sa: 2 Brooks 22703 (DS). Kit rae 5 mi E of Flagler, er pets S Brooks 2264 1 (DS). Logan Co.: E of Sterling, act 333 (MO, POM). 172 Co.: 5 mi S of Holyoke, Stephens & Brooks 24072 ids Prowers Co.: 20 mi S and 7 of Holly, Stephens & Brooks 21931 (DS). Sedgwick Co.: mi S of Julesburg, Bic tiene & & Brooks 24059 (DS). Yuma Co.: Wray, Eggleston in 1919 is , MO, POM). kansas: Barber Co.: 3 mi W W and 5 mi N of Medicine Lodge, McGregor 14430 ( KANU, SMU, US). Clark Co.: 8 mi S of Sitka, ix rg & Imler 766 (KANU, MO, NY). Ellis Co.: 2 mi W of Hays, T 97 ( ARIZ, CAN, KL, OKLA, PH, RM, SMU). Ellsworth Co.: 1 mi N and 1 of Kanapolis, dd dy Latham in 1950 (GH, KANU, TEX, US). Ford Co.: Ca. 5 mi ud of Dodge City, Towner 9 (DS). Grant Co.: High 1 ipland prairies, Thompson 1 (CAN, F, NY, UC, US). Harvey Co.: 8.5 mi E of Newton, Harms 1633 (SMU, UC). Kearney Co.: 5 mi E of Kendall, Rydberg & Imler 1059 (KANU, NY). Kingman Co.: 3 mi E of E Stephens 11127 (OKLA ). ME Co.: 11 mi E of Meade, Hubert 3593 (KANU, OKLA). Montgomery Co.: 2 mi S of Sycamore, McGregor 12839 (KANU, NY). Pottawatomie Co.: State Park No. 2, Marsh 1725 (KANU, SMU, US). Riley Co.: Stony hills, Norton 168 (GH, MO, NMC, NY, RM, US). 12 mi N of Manhattan, Raven & Gregory 19483 (DS). Scott Co.: 10.6 mi N of Scott EFA Towner 160 (DS). Smith Co.: 2 mi W of Cedar, Horr E131 (COLO, F, GH, KANU, LL, L., OKLA, RM, SMU, UC, US). Wilson Co.: 3 mi NW of Neodesha, McGregor 4306 (GH, KANU, US). Miussourt: Atchison Co.: No locality, Bush 10321 (GH, PH, POM). Iron Co.: Deo Arc, Smith 460 (F). OKLAHOMA: Alfalfa Co.: 3 mi N and 7.8 E of Cherokee, 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Stratton 6371 (OKL, OKLA). Atoka Co.: No locality, Hopkins et al. 1132 (OKL, RM, WTU ). Blaine Co.: Roman Nose State Park, Goodman & Waterfall 4189 (OKL, OKLA). 21.5 mi W of Kingfisher, Torte 153 (DS). Carter Co.: 4 mi N of Springer, Waterfall 705 ( OKL, OKLA, d mi S of Davis, Towner 144 (DS). Ca. 7 mi E of Fox, Towner 143 (DS). Cimarron Co.: 4 mi N of Kenton, Rogers 5697 (OKL, TEX, US). Comanche Co.: Boggy Hollow Creek, Eskew 1743 (OKL, OKLA). Dewey Co.: W of Vici, Goodman 2573 (GH, MO, NY, OKL, POM, RM, WTU). Ellis Co.: Near Shattuck, Clifton 31 55 (GH, NY, OKLA). Greer Co.: 2 mi S of Mangum, Robbins 3035 (OKL, SMU, UC). 6 mi S of Mangum, Towner 80 (DS). 2.7 mi S of Mangum, Towner 83 (DS). Harmon Co.: 13.5 mi W of Mangum, Waterfall 7766 (OKL, OKLA, TEX). Harper Co.: Near Buffalo, “hay 0 (DS, 3H, NY, OKL, OKLA, SMU). Kingfisher Co.: 8 mi E of e run 250 (K , OKL, UC). Lincoln Co.: 6.5 mi S of Perkins, Towner 148 (DS). 3.2 mi Se of Perkins bel 149a (DS). Logan Co.: wi 8 mi N of Guthrie, Towner 150 (DS). Major Co.: Togo, Demaree 12370 (MO, POM). 1.7 mi NE of Orienta, Raven & Gregory 19471 (DS). Murray Co.: Davis, ro. 12508 (G n ' MO, NY, OKL, PH). Turner Falls, Cory 19 25 (OKLA, SMU). 6.2 mi ) 3 N of Sulfur, Towner 146 (DS). 8.1 mi S of Davis, Towner 145 (DS). Oklahoma Co.: "s aw W of Woo od, Waterfall 2772 (GH, OKL, OKLA). Pontotoc Co.: 1.5 mi NE of Lawrence, Robbins 2995 (OKL, SMU, UC, WTU). Roger Mills Co.: Ca. 8 mi E of Strong City, Towner 155 (DS). Tulsa Co.: W of Tulsa, McKelvey 2508 (GH, POM). Woods Co.: Between Cimarron and Waynoka, CUR ss e Waterfall 4229 (COLO, KANU, OKL, OKLA). TEXAS: Anderson Co.: Palestine, Palmer 13426 (MO). Andrews Co.: 21-23 mi NE of l Correll. 32786 (LL). Aransas Co.: 4.6 mi NE of Rockport, McCart 5566 (TEX). Copano Bay, sandy beach, Bogusch S-75 (US). 1.3 mi W : Copano Village, 2 shore of Copano Bay, Towner 182 (DS). Armstrong Co.: Ca. 10 mi NE of Wayside, Rowell 5402a (OKLA). Au stin Co.: Colbert’s Station, P Sheldon 3575 (F). Bailey 5 55 "5 mi NW of Muleshoe, Correll a 13106 (LL, SMU). Brazoria Co.: 11.6 mi S of San Luis Pass Bridge on road to Surfside, Towner 173 (DS). Brazos Co.: College Station, Parks in 1946, 1947 (RSA, TEX). NW of Bryan, in prairie, Lundell & Lundell 11304 (POM, SMU). 3.5 mi S of College Station, McVaugh 6997 (F, LL, SMU). Calhoun Co.: Magnolia Be: ach, Tharp in 1930 (TEX). 0. d mi from shore at Magnolia Beach, Towner 178 (DS). Cameron Co. Stover Point, Laguna Atascosa National Vyas Refuge, 48 190 1125 (SMU, TEX). Boca Chica, Lundell & Lundell 9617 (DS, GH, NY, POM {U). Brownsville, Fisher 41188 (ARIZ, NEB). 0.8 mi N of bridge from ee on 9 js Towner 189 (DS). Childress Co.: 10 mi N of Childress, Correll & Johnston 16876 (LL). C ochran Co.: 14 mi N of Bronco, Toner 137 (DS). 10.3 mi N of Bronco, Towner 136 (DS). Collin Co.: Near Plano, Lundell & Lundell 9313 (DS, GH, LL, POM, SMU). Cooke Co.: 5 mi N of Gainesville, Gould 6867 (MO, SMU, TEX, UC). Dallam Co.: 1 mi SE of Texline, York & Rodgers 192 ( SMU, TEX, UC). Dallas Co.: Dallas, Reverchon 3563 (NY). Denton Co.: 5 mi N of Denton, Cory 57359 (SMU). Eastland Co.: Ranger, Robinson in 1931 (POM ). Bur Co.: Stephensville State Park, Hoisington in 1946 (TEX). Fannin Co.: Bonham, Milligan s.n. (NMC). Galveston Co. (?): saly eston I., Bechdolt in 1870 (PH). Gray Co.: McLean, Craig in 1934 OM). Grayson Co.: Near Gunter, van Meter 18 (SMU). Hale Co.: 7 mi S of Plainview, Gould 7156 (SMU). Hal Co.: Mab: Thames 7192 (TEX, US). Hardeman Co.: Acme, Russel 88 (TEX). Hartley Co.: 3 mi SE of Dalhart, Cory 32667 (POM). Hood Co.: Near Center Mills, Blackwell 26 (NY, SMU). um Co.: Borger, Hope 4 (LL). Jack Co.: 2.5 mi NE of Jacksboro, Hennen 421 (SMU). Jackson Co.: 11.2 mi W of Palacios, Towner 175 (DS). Johnson Co.: S of Rio Vista, 9 4 (SMU). Kleberg Co.: Beach along Laguna Madre, Laureles Division of King Ranch, Johnston 53224.13 (TEX). Padre I., Cory 49120 (GH, LL , SMU). Madison Co.: 3 mi N of North Zulch, Morgan 39 (TEX). Matagorda Co.: 5.5 mi S of Matagorda, Towner 174 (DS). Montague Co.: 4 mi N of Nocona, Whitehouse 10070 (SMU). Montes omery Co.: Willis, Warner s.n. (MO). Nueces Co.: Corpus Christi, Drushel 8932 (MO, NY, US). Corpus heen Heller 1517 (PH, US). Parker Co.: Weatherford, Tracy 7820 (F, GH, MO, NEB, NY, TEX, US). Parmer Co.: 7.8 mi NW of Farwell, Rowell 10023 (DS, RSA ). Refugio Co.: 4.1 mi RE of Austwell, Towner 181 (DS). Robertson Co.: 3-4 mi s: e Hearne, Reeves 930 (POM). San Patricio Co.: 2 mi S of ug n Cutler 920 (OKL, U). 3.5 mi S of Ingleside, ca. E mi from Corpus Christi Bay, Towne (DS Smith ds Troupe, Reverchon 2744 (MO, US). Tarrant Co.: Sandy soils, Pd. Mon 913 (F, GH, KANU, NEB, NY, PH, UC). oe Zandt Co.: 3 mi E of Wills Point, Shinners 12381 (COLO, SMU). Walker Co.: Vicinity of Huntsville, Dixon 569 (F, POM, RM, US). Wise Co.: Near Park Springs, McCart 1633 (SMU, TEX). New Mexico: Chaves Co.: `8. 3 mi W of Caprock, Towner 134 (DS). De Baca Co.: 11 mi S of Ft. Sumner, Towner 131 (DS). Eddy — — 1977] TOW NER—CALYLOPHUS 115 Co.: Lakewood, Wooton in 1909 (NMC, US). Guadalupe Co.: dad between Anton Chico and Santa Rosa, Arsène & Benedict 16681 (POM, US). Lea Co. nowles, 2: in 1909 (NMC, US). Roosevelt Co.: 5 mi NE of Portales, Goodman & Do 1119 (DS, F GH, MO, NY, PH, POM, RM, UC). 14.0 mi SW of Elida, Towner 133 (DS). 8.0 mi E of Tai iban. Towner 132 (DS). San Miguel Co.: Between Las Vegas and Romeroville, Arsène d» Be nedict 15458 (POM, US). Union Co.: Perico, Bartlett 227 (NMC). arizona: Graham Co.: Willow Spring, 5 481 (GH, US). Navajo Co.: 4 mi N of Carrizo, Pulta b Phillips 1008 (ARIZ, UC). 4 mi SW of Show Low, Lehto 1072 (ARIZ). Forestdale, 66 mi S of —— EUN S3 (US). O. CHIHUAHUA: 18 mi N of Rubio, District of Cusihuiriachic, Shreve 7960 (POM, US). rAMAULIPAS: Coastal n near Río Grande, Le Sueur 328 (probably of this species, ARIZ, F, TEX). This species, the earliest described, cultivated, and illustrated ( Hooker, 1825) taxon of Calylophus, occurs widely over the North American Plains, and is the most familiar member of the genus, although its breeding system has only re- cently been described (Towner, 1970b). Permanent translocation heterozy- gositv, half-sterility of pollen and ovules, self-compatibility, and small flowers enhancing self-fertilization are present in C. serrulatus as part of the genetic system of complex structural heterozygosity. This type of breeding system is frequent in the Onagraceae, but in Calylophus is restricted to this species. Charac- ters associated with this breeding pattern are used as the primary basis for dis- tinguishing C. serrulatus from C. berlandieri. Whether factor complexes exist and are maintained in C. serrulatus by ga- metic lethals was not determined. The presence of gametophytic half-sterility makes the involvement of gametic lethals appear likely, but does not exclude mechanisms utilizing megaspore competition and/or zygotic lethals. Although unlikely, the observed levels of pollen and ovule sterility may stem from random chromosome disjunction and the resulting genetic deficiencies and duplications in the gametes. There were no consistent reciprocal differences in pollen fer- tility, morphology, or chromosomal configurations in hybrids between C. ser- rulatus and C. berlandieri, results which would be expected if gametic lethals regulated the transmission of factor complexes. A system utilizing self-sterility alleles combined with egg lethals (see Steiner, 1956, 1957) does not seem to be operating in C. serrulatus. All reciprocal crosses between C. serrulatus and the self-incompatible species C. berlandieri produced only self-compatible hybrids, whereas some self-sterile progeny would be predicted if C. serrulatus had re- tained a functional self-sterility allele. The flowers of C. serrulatus are generally identical to those of C. berlandieri except for their smaller size, relatively shorter filaments and style, and the po- sition of the stigma. The placement of the stigma among or near the anthers and early dehiscence of the anthers frequently causes flowers to self-pollinate before anthesis. Undisturbed flowers in the greenhouse showed a high level of autogamous seed set. The similarity of the flowers of the two species extends to morning anthesis times and the ultraviolet-absorbing areas on the petals, stigma, and stamens. Morning anthesis had been known from the first observa- tion of the species by Nuttall (in Hooker, 1825), and was seen and reported again by Stevens (1920), but this was apparently not known to Munz (1965) or Raven (1964). Both mentioned only vespertine anthesis for Calylophus. Insects do not 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 seem to visit C. serrulatus frequently. Stevens (1920) observed that bees ignored C. serrulatus, although they were active on nearby plants of Gaura and Oenothera. In my own collecting at 31 colonies of C. serrulatus, done at all times of day, no insects were observed visiting flowers. Halictid bees (Agapostemon, Evylaeus, Dialictus) were observed during the morning at one population in Major Co., Oklahoma (P. Raven, personal communication), however. Meiotic configurations of 34 individuals from 28 populations consisted of a definite or probable ring or chain of 14 chromosomes. Five plants had a ring or chain of 12 chromosomes plus 1 bivalent. Seventeen plants from 5 popula- tions were examined and found to be self-compatible. In Calylophus the corre- lation of self-compatibility and complex structural heterozygosity was found to be nearly perfect. Two exceptions were individuals of C. berlandieri having a ring of 12 plus a bivalent at meiotic metaphase With a broad range of phenetic variation which largely overlaps that of C. berlandieri, C. serrulatus cannot be reliably diagnosed without the use of floral characters or pollen fertility. Parallel geographical variation in vegetative parts is such that near most areas of contact, the two species are unn similar. This implies that C. serrulatus, almost certainly a derivative of C. berlandieri, is of multiple origins, has experienced secondary local intr mc nis the parental species or has responded in a similar fashion to natural selection. Much of this vegetative variation in C. serrulatus follows a smooth east-west cline, whereas the variation is more discontinuous in C. berlandieri. Thus the discontinuities in the parent species are not reflected in the derivative. Populations formerly assigned to C. australis, because they closely resemble adjacent populations of C. berlandieri, and by their relative geographical sepa- ration from the bulk of C. serrulatus, may be an exception to the introgression hypothesis. They could well have been independently derived from C. ber- landieri. In this paper they are synonymized because no firm knowledge is avail- able on the phylogeny of other populations of C. serrulatus. Thus, to preferentially recognize C. australis, with scant genetic or morphological support, is to ignore possible polyphyly in the rest of C. serrulatus. Instead, C. serrulatus is best recog- nized as a complex assemblage of populations having a common breeding sys- tem. These populations encompass a broad morphological diversity, and some of them may have been evolved separately from the bulk of the species. Variation in C. serrulatus involves leaf size and shape, stature, pubescence, and flower size. Flower size is variable throughout the geographical range. Some of the largest-flowered forms occur near large-flowered populations of C. ber- landieri subsp. pinifolius in central Oklahoma. Although the vegetative charac- ters are clinally distributed, most populations occurring west of approximately 98°W longitude are comprised of well-branched, short-leaved, and relatively low-statured plants. East of that line plants are generally less branched, taller and more erect, long leaved, and more densely strigose-canescent. A few representatives of the eastern form are the following: Davis, Murray , Oklahoma, Demaree 12508 ( GH, MO, NY, OKL, PH). Elmore, Farribault Co., Minnesota, Pammel 596 (GH, MO, RM, US). Near Plano, Collin Co., Texas, Lundell & Lundell 9313 (DS, GH, LL, POM, SMU). Examples of the western 1977] TOWNER—CALYLOPHUS 117 phenotypes are: 1 mi S of Texline, Dallam Co., Texas, York & Rogers 192 (SMU, TEX, UC). Wolhurst, Douglas Co., Colorado, Clokey 3827 (CAN, DS, F, GH, MO, NY, PH, POM, RM, SMU, UC, US, WTU). Moose Jaw, Saskatchewan, Turner 48 (GH, NY, POM, RSA). Representatives of the coastal Texas popula- tions formerly assigned to C. australis include all specimens cited above from Brazos, Austin, Galveston, Brazoria, Matagorda, Jackson, Calhoun, Refugio, Aransas, San Patricio, Nueces, Kleberg and Cameron cos. Observed instances of direct contact between C. serrulatus and C. berlandieri were rare. No hybrid zones or populations were identified unequivocally, and the two species were locally allopatric. The only evidence obtained concerning possible mixed populations or direct contact came from the following collections: 3.2 mi S of Perkins, Lincoln Co., Oklahoma, Towner 149 (DS), one short-styled plant seen in a population of long-styled C. berlandieri subsp. pinifolius, one of which had a ring of 12 chromosomes and one pair at meiosis. 11.1 mi S of Perry- ton, Ochiltree Co., Texas, Towner 158 (DS), chromosome counts of 3 pairs + ring of 4, and possible ring of 12 + 1 pair from the same population, thus perhaps some individuals of C. serrulatus in a population of C. berlandieri subsp. ber- landieri. 11.2 mi W of Palacios, Jackson Co., Texas, Towner 175 (DS), this was a population of C. serrulatus which was only 2.2 mi E of a colony of C. berlandieri subsp. berlandieri ( Towner 176), the two being identical except for pollen counts and the longer style lengths in the latter population. Sympatry of C. serrulatus with members of sect. Salpingia is frequent. In the southern Great Plains, C. hartwegii subsp. fendleri is often found near or adjacent to populations of this species. Less commonly, C. hartwegii subsp. pubescens and C. lavandulifolius occur with C. serrulatus in the same region and at the eastern base of the Rocky Mountains. Lastly, in eastern and southeastern New Mexico, C. hartwegii subsp. filifolius occasionally comes in contact with C. ser- rulatus on calcareous plains east of the Pecos River. In none of these cases has any evidence of introgression or hybridization been observed. LITERATURE CITED CLELAND, R. E. 1951. Extra, diminutive chromosomes in Oenothera. Evolution 5: 165-176. — 967. Further evidence bearing upon the origin of extra diminutive chromosomes in Oenothera hookeri. Evolution 21: 341-344 B. HYDE 963. Evidence of relationship between extra diminutive chromo- somes in geographically remote races of Oenothera. Amer. J. Bot. 50: 179-185. CockERELL, T. D. A. 1903. Notes on the entomology of Pecos, New Mexico. Canad. Entomol. 35; 342-343. sg de W. A. & P. H. Rave 1973. Distribution of the chalcone, isosalipurposide, in -808 e Onagraceae. Bomb 12: 80 — —— — 1974. Pigments e for ultraviolet patterns in flowers of Oenothera (Onagraceae). Nature 252: 705-706. eed S. 1840. Genera Plantarum Secundum Ordines Naturales Disposita. Fr. Beck Universitatis Bibliopolam, Wien. [1836-1840 for the whole wor Grecory, D. P. 1964. Hawkmoth pollination in the genus ences (part 2). Aliso 5: 385-419 —— —— & W. M. Krem. 1960. Investigations of meiotic chromosomes of six genera in the 5 Aliso 4: 505-521 HacEN, C. W., Jr. E A 0 to the cytogenetics E the genus Oenothera with spe ecial reference to certain forms from South America. In R. Cleland (editor), Studies in Oenothera C iosenetids and Phylogeny. Indiana Univ. Publ. Sci. Ser. 16: 305-348 118 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 HÁkANssON, A. 1942. Zytologische Studien an Rassen und Rassenbastarden von Godetia whitneyi und verwandten Arten. Acta Univ. Lund, n.s., 38: 1- P Supernumerary chromosomes in Godetia viminea. Hereditas 35: 375-389. Hooker, W. 1825. Oenothera serrulata. Exotic Flora 2: pl. 140. KURABAYASHI, M, H. Lewis & P. H. Raven. 1962. A comparative study of mitosis in the nagraceae. Amer. J. Bot. 49: 1003-1026. Lewis, H. 1951. The origin of supernumerary chromosomes in natural populations of Clarkia elegans. Evolution 5: 142-157. 1 The mechanism of evolution in the genus Clarkia. Evolution 7: 1-20 195 . H. Raven, C. S. VENKATESH & H. L. WEDBERG. 8. Observations of piik Finn in the pee Aliso 4: 73-86. Loney, & J. Brux. 1957. L'incompatibilité dans Oenothera serrulata Nutt. Experientia —24, MAzOKHIN-PORSHNYAKOV, G.A. 1969. Insect Vision. Plenum Press, New York. 306 McKe vey, S. D. 1955. Botanical Exploration of the Trans- -H West 1790. "1850. Arnold Arboretum of Harvar Univ., Jamaica Plain, Massac Munz, P. A. 1929. Studies in Onagraceae IV. A revision of the i Salpingia and Calylophis of the genus Oenothera. Amer. J. Bot. 16: 702- Onagraceae. N. Amer. Fl., ser. 2, 5: 1-278. ÖSTERGREN, G. 1947. Heterochromatic B-chromosomes in Anthoxanthum. Hereditas 33: —296. RAFINESQUE, C. 1819. The genera of North American plants and a catalogue of the species to un year 1817. By Thomas Nuttall. (Review.) Amer. Monthly Mag. & Crit. 196 ev. 4: : Raven, P. H. 1962. The systematics of Oenothera subgenus Chylismia. Univ. Calif. Publ. ot. 34: 1-122. 1964. The generic subdivision of Onagraceae, tribe Onagreae. Brittonia 16: 5 & D. P. Grecory. 1972 a. A revision of the genus Gaura (Onagraceae). Mem. Torrey Bot. Club 23: 1-96. —. 1972b. Observations of meiotic chromosomes in Gaura (Onagraceae). 5 24: 71-86. SHINNERS, L. H. 1964. Calylophus (Oenothera in part: Onagraceae) in Texas. Sida 1: 337—354 Spach, E. 1835a. Onagraíres. Histoire Naturelle des Vegétaux. Phanérogames 4: ee . 1835b. Monographia Onagrearum. Nouv. Ann. Mus. Hist. Nat. 4: 320-4 STEINER, E. 1956. New aspects of the balanced lethal mechanism in Oenothera. p tics 57. Further a eg of an incompatibility allele system in the complex-hetero- zygotes of Oenot r. J. Bot. 44: O. A. 19 0. Eum on species of DN visiting evening flowers (Hym.). 4. TowNEn, H. 1970a. Calylophus. Pp. 1121-1123, in D. S. Correll & M. C. Johnston, Manual of the Vascular Plants of Texas. Texas Research Foundation, Renner, Texas. 1970b. Evolution and breeding systems in Calylophus (Onagraceae). Ph.D. dissertation, Stanford Univ., Stanford, Calilam ia. AVEN. 1970. A m species and some new combinations in Calylophus 245. (Onagraceae). Madrono 20: 241- INDEX OF LATIN NAMES Numbers in bold face refer to descriptions or names listed in synonymy; nan are recognized as valid; names in italic refer to synonyms; numbers with asteris to names incidentally mentioned. Acacia 76* Acmaeodera 110* Agapostemon 74*, 77*, 87*, 91*, 95*. 106*, 110*, 11 Anthophora 106* Anthoxanthum 59“ Apis 74*, )* Artemis 5 88* Asteraceae 62* Atalopedes campestris 110* Augochlorella 110* Bombus 77*, 87*, 110* Buprestidae 110* — 49* — 48* 59», 59*-65*, 67, 87*, 91*, 97*. 99, 115*-117 Bo 5 48 * 50*, 65*- 67* 68*. 95 * C . 09 * —sect. e pia , oo", 59*, BIS. 65*— 68, 70*, 75*, 77, 81*, 86*, po 91*, 93*. 95*. 96*. 97*. 106*, 110% 117* australis 50* 111, 116* m E 50*—51*. 61*—62*. 66*, 68* 81*, 87*, 96*, 100, 102*, 105*. EA 1115 115*-117* — subsp. berlandieri 51*—52*, 59*, 60*, 65* 78*, 97*. 1OL*, A 103, 1051065 108-109 *, 117 — subsp. pínifolins 32 5960 63 * 101*—102*, 105-106 * 107, 108*— 111*, 116*-117* drummondiana 110 drummondianus 48*, 50*, 102*, 103, 108* subsp. berlandieri 1033 —subsp. drummondianus 107, 110 hartwegii 48*, 50*—51*, 59*, 61*—62*, 65*— 67*, 68, 69*—70*. 74*, 8788, 91*— 96*, 99*. 102 * EN fendleri 50*, 53*, 60*, 64*, 66*, [ 74*, 80*, 81, 83*, 8 91 97 *_98*, 106* =< filifolius 50*—51*, 54* 60*, 65*, 74*-75* 77*, 18, 79817 83*, ue 98*. 106*, 110*, 117* n : artwegii 50*-51*, 54*, 56*, 0 * —66*, 70, 71, 72“, 74*— T7. 79*, 81*, 8687 91 *, 99 * — lavandulifolius 88 —subsp. maccartii 50*-51*, 55*, 60*, 65*, 70*-72*, 74*, 7 81*, 106*, 110* —subsp. pubescens 51*, 5 yes in roman refer ks (* 5, 77*-78*, 86*-87* 91" 98*. 106*, 110*, 117* —subsp. foumess 92 ; ) —var. lavandulacfotius 88 — var. maccartii . toumeyi s 78 —var. hartwegii 71, 74*, 78, 81 —var 92 lavi andulifolius 48* 50*. 56*, 59*—61*, 65*, 7*—70*, 75*, , 87*, 88, 89*. 91“, 98* 102*. 106*. 117* nuttallii 67 *, 110 serrulatus 48*, 50*-51*, 57*, 59*-63*, 66*— 63", 70%, Si, 84* B7*, 102*, 103*, 108*, 110, I —subsp. serrulatus 103 — var. spinulosus 107, 110 ee 48*, 50*, 58*, 60*—65*, : 89*. 92. 93*, BI*. 100*, 112*. 115*-— *. 68*, 69*— bien pr 59*. 61*, G5* —66*. 68*. 70*, 94, 95*—99*, 102*. 106* tite sido 50*, 58* —97*, 98. 9f irt A a 58*, 91* 60* 63% 95*, 96, 97*, 99*. 110* C amissonia pes Ce opm 61*, 70* Cercocarpus 88* Clarkia 49*, 59* amoena 51* —subsp. amoena 51* unguiculata 51* Dialictus 87*, 95*. 110*, 116* Evylaeus 77*, 83*, 87*, 95*, 106*, galpinsia 83* Galpinsia wa 67, 68 camporu 4 1 qhana 96, 98* fe EN 2 filifoli Eee 96 greggii hartwegi —var. fendleri 81 eai 67*, 71 id”, * 110, 116* 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. interior 84 lampasana 84 slag wa a ar. dde 88 Gaura 49*, 51*, 59*, 65*, 116* Gayophytum 59* Halictus 110* Hy les cee 61*, 69*, 87*, 91*, 95*, 106* Hauya 49* Heterogaura 4 49* Juniperus 78*, 82*, 85*, 88* Larr ea divaricata 76*, 78*, 88*, 96* Lasioglossum 93 Manduca 91 quinquemaculata 87* capillifolia 107 drummondiana 110 i 107, 109* intermedia 111 melanoglottis 107, 109* oblanceolata 111 serrulata 67*, 110 £e a eo = RU a Š Š š — — var. drummondii 110 var. nuttallii 110 var. pinifolia 107 r. spinulosa 110 PA A 110 EUN 48*, 59*, 62*, 66* —tribe Onagreae 48*, 49* 98 4851“, 59“, 116* —subg. Salpingia 49*, 67*, 68 —subg. Calylophus 99 ens —var. pringlei 71*, 74*, 75 pica 84 hartwegi var. fendler —subvar. 1 88 — var. tubicula 96 di 68, 79*, ^ — var. fen ndleri e asa hee 78 8 —var. hartwegii 71, 78, oes 83* var. lavandulaefolia 8 var. 1 92 var. a 71, 74*, 78, 81, 88 5 s lamp dae, 67*, 74* 88 r. glandulosa 88, 91* typica 71 „ . 71, 74*, 88 ei pringlei 71, "e serrulata 105*, 110 —subsp. drummondii 103, 107, 108“, 110 —subsp. pinifolia 3 105*, 107 —subsp. serrt 1 110 var. douglasii var. 5 103, 107, 110 —f, flav 1 —var. integrifolia 111 —var. maculata 10 var. nuttallii 110 —var. pinifolia 103, 105*, 107 var. 170 ilosa 110 a 103, 110 x X serrulatoides 96 —var. demissa 96 —var. d 78 5 76*, Pinu eae 82* monophylla 88 ponderosa 82*, 88* Prosopis 103* glandulosa 76*, 82*, 85* 0* 1 7 179. 96* CHROMOSOME NUMBERS IN LEGUMES'! PETER GorpBLATT" AND G. DAvipsE® ABSTRACT Chromosome numbers are reported for 39 species representing 35 E pa The lowing are new generic (and specific) reports: Acrocarpus fraxinifolius 2n — 24, Ambla ee andongensis 2n — 28, Baikiaea d 2n = 24, Brasilletia mollis 5 tschya ae schynomenoides 2n = 40, K. africana 2n = 30, K. strigosa 2n = ca. 36, Maac ckia amurensis 2n 18)20, Paramacrolobium nine ial um 2n — 24, Schizolobium d 2m = 26, Sphaen ion salsula 2n = 16, Sphenostylis marginata 2n = 22, Strongylodon macro- botrys 2n = 28, Tachigalia paniculata 2n = 26, Tylosema fassoglensis 2n — 52, Xanthocercis zambeziaca 2n = 26, and Virgilia oroboides 2n = ca. 54. New species reports are the following: Cordyla africana 2n = 20, Desmodium barclayi 2n. = 22, Dioclea virgata 2n. = 22, Entada pursaetha An = 28, Erythrina livingstoneana 2n = 42, Mucuna sloanei 2n = 22, and Sindora wallichii 2n = 24. This paper is preliminary to a general review (P. H. Raven & P. Goldblatt, in preparation) of cytology in the Leguminosae. This work is being undertaken in conjunction with other systematic research in the family to be presented at the Legume Conference planned for 1978. The majority of counts reported here are for species and genera selected to fill some of the many gaps in the cytology of the family. Although there are a large number of chromosome counts in the Leguminosae, the family can still be said to be in general poorly known cy- tologically with an estimated 18% of the ca. 18,000 species having been investi- gated (Bandel, 1975). The present paper idadi; chromosome counts for 39 species in 35 genera. Of these, 15 are first reports for genera. MATERIALS AND METHODS Root tips only were used in this study, hence all counts are mitotic. Root tips were pretreated either in cold water for 24 hours and stained using the Feul- gen technique or were placed in 0.003 M hydroxyquinoline for 4-5 hours and stained in lactopropionic orcein. Species studied are listed in Table 1 and al- most all plants have herbarium vouchers which are housed mainly at Missouri Botanical Garden (MO), but also at New York Botanical Garden (NY) or else- where. DISCUSSION The results are listed in Table 1 and are discussed by subfamily and tribe. The three traditional subfamilies Papilionoideae, Caesalpinioideae, and Mimo- "We would like to thank Dr. B. A. Krukoff for his energetic and untiring efforts to b ain of Leguminosae. Almost all wu used in this study were obtained either directly or 0 through him. We would also like to thank the various individuals and 5 too numerous to me 1115 on who have provided us with material for study. Kru ikoff Curator of 14 Botany, Missouri Botanical Garde m, 2345 Tower Grove VAE h St. Louis, Missouri 631 Botany Department, ME Botanical Garden, 2345 Tower Grove Avenue, St. Louis Missouri 63110 ANN. Missount Bor. Garp. 64: 121-128. 1977. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 punt 1 bo soideae are recognized. Tribal subdivisions are those of Hutchinson (1964) for Mimosoideae and Papilionoideae, while Taubert’s (1891-1894) treatment is used for Caesalpinioideae as presented recently by Heywood (1971). Previously published chromosome numbers referred to in this paper are taken from the standard chromosome indices, particularly Darlington & Wylie (1955), Fedorov (1969), and the annual Index to Plant Chromosome Numbers, the most recent of which summarizes reports for 1972 (Moore, 1974). MIMOSOIDEAE Adenanthereae.—The report of 2n = 28 in Amblygonocarpus is a new generic record. This count accords with n = 14 in Prosopis and other allied genera in the tribe where x = 14 is probably basic. Low base numbers of x = 13 in Adenanthera, Piptadenia, Neptunia, and others, and x = 12 in Calpocalyx and Xylia are most likely derived. The count for Entada purseatha is a new species record and in accord with n = 14 reported for several other species of this genus. Ingeae.—The count of 2n = 26 for Samanea saman confirms a previous record for this species. A base number of x — 13 has been reported in several genera of the tribe with the only exception, Calliandra, x — 8, standing out here. Counts for more genera in this tribe are needed before the significance of the exceptional x — 8 can be assessed. CAESALPINIOIDEAE Caesalpinieae —Counts in this group for Caesalpinia pulcherrima (2n = 24) and Delonix regia (2n = 28) confirm previous reports for these species. Counts for Acrocarpus (2n = 24), Schizolobium (2n S 26), and Brasilettia (2n = 24) are new generic records. With x — 14 probably basic in this alliance and with x — 12 and 11 well established in several other genera, present counts are con- sistent with the cytological pattern of this tribe. The number 2n — 26 in Schizolo- bium is, however, a new number for Caesalpinieae. Detarieae.—The counts for Baikiaea, 2n = 24, and Tachigalia, 2n = 26, are new generic records. The number in Baikiaea is consistent with the majority of chromosome numbers in Detarieae where x — 12 is probably basic. Other num- bers in the tribe are n — 10 and n — 8 reported in only a few 1 0 The record of 2n = 26 for Tachigalia paniculata is a new number for the tri The count for Sindora wallichii is a first report for this species. Other counts in Sindora are n = 12 in S. siamensis, n = 10-12 in S. cochinchinensis, and n = 8 in S. supa. Unless Sindora is cytologically heteroploid, n = 12 would seem most likely correct for the genus. Sindora supa should, however, be checked to verify the unusual n = 8 reported by Atchison (1951) The count of 2n = 24 for Afzelia africana confirms previous reports for this species. Amherstieae The report for Paramacrolobium is a new generic record and the number 2n — 24 is consistent with x — 12 in the seven other genera of Am- herstieae known cytologically. Cerceae.—The count for Tylosema fassoglensis, a segregate of Bauhinia ( al- 123 SGUMES IN LE '"MBERS CHROMOSOME D H GOLDBLATT & DAVIDSI 1977] (OW) VFSS6 YANDI VUND WU VA UYON “LArypeyy cc WOH Y IOL (AWOSJON ) sisuə]Baosspf muəso]ñ J , , '(OIN) 9669 ?spiap(qp "VH Woy wu siq Poria, vuy FZ PROYpyy-eupY (ummyos ) nsuimuot] Duzijsoitg 3922197) COW) 6I&/FL uapp Q uəpD, 'jso10,| epng “YSA APAY ‘“eAuoy Tc pieuooT (CNEL) unopnao02. wun1q00]049DUDADQ , x ORATISIOYULY '(OIN) Se fiiuoz) ‘ueq jap Lueng ‘ooy ju) 9G quy. 23njnorupd. DIDTIYID [ y + (OW) 268IVS powyy ‘powan ‘upe Srurcjog o10dezutq F€ "uog %% / v ni0putS, (AN) “Us 4/0 SHOT “LOAY e[ounipunpurury apəu `qsiq Əuedn ‘visapoyy yc SULH Dpznlunjd. DODINIDS y x '(OIN) FS jo 12 uopp, Vuy ‘uoy ustoysey `ruruo PZ quis pupoujp pi]əz fv ovaLiejo(] (AN) FS Sid PNW Upg ^eieq (zeig 9c APTA CIPA) "nqfipapd. 1niqojozi2g, » (OW) 62 F2 9 P upo vyouoy ‘uoy uiojsezp “pupu Sc ‘FEY (` ool) visas xiuoJaqq "(OIN ) (Ws UOSJAN `XNIO4O gs "spue[s urg rc ZHENG CI) Duttayajnd pruidjpsavy (OW) AGFF 2sprap([ 'enoseq v| op ope y ieou toonen "e[snzouoA PG dy x uogrq (`N H) $1]]JOUL DIZIAIISDLG x x FISI/FZ p 49 uopp4 c(poyean[no) ^eNB[[ ^as nquerıy ^PXuoN F€ "UlV S QUSTAA Snqofiuixpif SNAIDIOLIY y x aeorurd[esor?) AVACIOINIdD TWSAVO `C ON) O F "opp Q uopp, `joumnpy 'uoməyw Ulojsv;[ "vuv 9c “Moy (Poef) unus paunung ovosu '(OIN) CYA D 22 uoppq [i] OvURAN un waseq vuv) ra ‘OC ?opsand. ppp, '(49Q9no4 OU) “U's uop407) “UOT}RIS SJEA OO ^rsopouy 9c PHOT S PXA CAYO Xo “AOA ) sieuozuopun sndap20uozfi]quiy » , 9vo19tgueuop y AV HCOIOSOINIIN uomnpuriju] 19 ono4 pue uo uc soroodg loq] tun N 9UIOSOLIO.U7 “snuos yey} 10) q310do1 ISI} ƏY} ƏV (,,) SNSLI9]SP OM] Aq pepe.01d sərnuə :sorods yey) 10] ο,,.e sa ƏY} aie (, ) yslio}se 9uo e 132 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 curling in drying; faucal annulus only slightly raised and somewhat fleshy, nearly obscured by the corona; corona remotely 5-gonal, fleshy, the margins thickened, rugose; ovaries 3-ridged, glabrous. Follicles unknown. This species is typically found at elevations from 1,200 to 1,800 m in Costa Rica, Honduras, Nicaragua, and possibly Guatemala. A distinctive species, Gonolobus rothschuhii is separated from all other Pana- manian members of the genus by its very narrow corolla lobes and color, and by the shape of its anther appendages. JARIEN: Cerro Tacarcuna, south slope, ridge-top forest well below summit, premontane wet b 1,250-1,450 m, 26 Jan. 1975, Gentry & Mori 13921 (MO). LITERATURE CITED SPELLMAN, D. L. 1975. Asclepiadaceae. In R. E. Woodson, Jr. & R. W. Schery, Flora of Panama. Ann. Missouri Bot. Gard. 62: 103-156. NEW SPECIES OF GIBSONIOTHAMNUS (SCROPHULARIACEAE/BIGNONIACEAE) AND TOURNEFORTIA (BORAGINACEAE) FROM EASTERN PANAMA AND THE CHOCO! ALWYN H. GENTRY? ABSTRACT new s are described from wet forest regions of eastern Panama and the Chocó of Colombia e ben alatus A. Gentry, Gibsoniothamnus mirificus A. Gentry, and Tournefortia tacarcunensis A. Gentry & Nowicke. Gibsoniothamnus alatus A. Gentry, sp. nov. Frutex epiphyticus. Ramuli irregulariter teretes. Folia elliptica, acuta vel acuminata, cuneata, glabra praeter domatia ciliata. Flores singulares, i un glabris. Calyx late alatus, ad instar stellae, alis ultra 1 cm onis. Corolla ( non vidi) a Epiphytic shrub. Branchlets irregularly terete to subangulate, very sparsely pilose. Leaves elliptic, acute to acuminate, cuneate at the base, chartaceous to subcoriaceous, glabrous above and below except for ciliate domatia in the axils of the lower secondary nerves, gland dotted below, the margin entire, very slightly or not at all revolute, drying dark olive above, light olive below, the secondary veins plane or slightly impressed above, prominulous to prominent below. In- florescence a single flower; pedicel glabrous, 1.5-2 cm long. Calyx very broadly winged, glabrous except a few trichomes near the ends of the wings, almost star shaped, ca. 6-7 mm long and wide without the wings, the wings each over 1 cm long, tapering to an acute point. Corolla (not seen) white. Pistil 23 mm long, the ovary globose, the style slender, 18 mm long. Fruit white, covered by the calyx. TYPE: PANAMA. DARIEN: N slopes of Cerro Pirre, lower montane rain forest (cloud forest), 700-950 m, 6 Apr. 1975, Mori & Kallunki 5449 (MO, holotype; isotypes to be distributed ). DS Additional collection examined: PANAMA. DARIEN: Cerro Campamento, S of Cerro Pirre, elfin forest, 20-22 Mar. 1968, Duke 15657 (MO). This species is utterly distinct in the genus because of its laterally winged, star-shaped calyx. Its closest relative is G. pterocalyx A. Gentry but that species has much narrower longitudinally oriented teeth. Gibsoniothamnus mirificus A. Gentry, sp. nov. Frutex epiphyticus. Ramuli irregulariter subangulati, pilosi. Folia obovato-elliptica, obtusa, cuneata, conspicue pilosa. Flores singulares, pedicellis pilosis. Calyx cupulatus, pilosus, valde 5-dentatus, dentibus linearibus, 2-2.5 cm longis. Corolla tubulosa, rubra. * Support for this work was provided by NSF grant OIP75-18202. ° Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. ANN. Missovni Bor. Garp. 64: 133-135. 1977. 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Epiphytic shrub. Branchlets pilose, irregularly subangulate. Leaves obovate- elliptic, obtuse to acutish at the apex, cuneate at the base, chartaceous to sub- coriaceous, 3-9 cm long, 1.5-4 cm wide, conspicuously pilose with 1-2-mm-long trichomes above and below, the margin entire, very slightly or not at all revolute, drying brownish olive above, tannish yellow below, the secondary veins plane above, prominent below; petiole densely pilose, ca. 5 mm long. Inflorescence a single flower; pedicel conspicuously pilose, 0.7-1 cm long. Calyx cupular, pilose strikingly 5-toothed, 5-7 mm long and 4-6 mm wide without the teeth, the 5 linear teeth exceeding the calyx by 2-2.5 cm, pilose, extended along the calyx as lateral ridges. Corolla red, tubular, glabrous, 3.5-4.3 cm long and 0.5-0.6 cm wide, lobes 3 mm long, with ciliate margins. Fruits not seen. Type: PANAMA. COLÓN: Santa Rita Ridge Road along trail from end of road (10.6 km from Transisthmian Highway, 3 km beyond hydrographic station) to Río Indio, 380 m, 13 Apr. 1976, Croat 34298 (MO, holotype; PMA, isotype). Additional collection examined: PAN LÓN: Plant collected by H. Wiehler on Santa Rita Ridge, cultivated at Marie Selby ines Gardens, Sarasota, Florida, Dressler MO). The striking calyx teeth of this species are by far the longest in the genus. It is otherwise similar to Costa Rican G. epiphyticus (Standl.) L. Wms. which is also more or less pilose throughout but has very much shorter calyx teeth, a fasciculate several-flowered inflorescence, and more coriaceous leaves. Tournefortia tacarcunensis A. Gentry & Nowicke, sp. nov. Herba erecta. Folia anguste elliptica, acuta, cuneata, subsessilia, glabrescentia. Inflores- centia scorpioidea, floribus sepalis lanceolatis, ca. 4 mm longis, corollae tubo 8-9 mm longo, lobis ca. 1.5 mm longis. Herb 0.2-0.5 m; stems glabrescent. Leaves alternate, narrowly elliptic, acute, cuneate at the base, entire, with 4—7 pairs of strongly ascending secondary nerves, 8-28 cm long, 2-6.5 cm wide, glabrous above, glabrescent below, rather suc- culent when fresh, membranous when dry, drying dark brown above, tannish gray below; petiole essentially lacking. Inflorescence scorpioid, contracted, 3-4 cm long, terminal; pedicels mostly 1-2 mm long. Calyx of 5(6) free sepals, lanceo- late, ca. 4 mm long, sparsely puberulous; corolla orangish to greenish cream, sparsely puberulous outside, the tube 8-9 mm long, the lobes ca. 1.5 mm long; stamens 5(6), borne near the apex of the corolla tube, sessile, the anthers ca. 1.2 mm long; ovary ovoid, the style ca. 6 mm long, the stigma conical, 1 mm long. Fruit not seen. TYPE: PANAMA. DARIÉN: Cerro Tacarcuna, W ridge, trail from summit camp to waterfall E of camp, 1,550-1,700 m, lower montane wet forest life zone, herb 0.5 m, flowers greenish cream, turning tannish, inflorescence scorpioid, stamens equalling petal number, ovary 2-locular with axile placentation, 2 Feb. Gentry & Mori 14114 (MO, holotype). > Additional collection examined: CoLomBia. cHocd: Slopes of Serrania del Darién, E of Unguia, premontane wet forest, ca. 1,300 m, herb 0.2 m, flowers orangish, 19 July 1976, Gentry, León & Forero 16772 (COL, MO). 1977 SPELLMAN--PANAMANIAN ASCLEPIADACEAE 135 This species seems remarkably distinct from all other members of the genus. It is apparently the only clearly herbaceous species of Tournefortia (a com- pletely unrelated species, T. sibirica L. is a wiry herb but often segregated as Messerschmidia). Another unusual feature is a tendency to 6-parted flowers. The pollen of T. tacarcunensis is of the type described as “Type IL," 3-col- porate, subprolate with expanded poles, psilate at the poles and verrucate at the equator by Nowicke & Skvarla ( 1974) The closest relative of T. tacarcunensis may be T. ramonensis Standl. of up- land Costa Rica and Chiriqui Province—though omitted from the Flora of Pan- ama treatment of the family ( Nowicke, 1969). That species has generally similar flowers which differ in the corolla being densely and rather strigosely pubescent outside. The inflorescence of T. ramonensis is also much more elongate and the flowers are essentially sessile. Vegetatively T. tacarcunensis differs con- spicuously in its glabrescent, narrowly elliptic leaves which are narrowly cuneate, essentially sessile, and have only about 6 pairs of secondary nerves. Another Costa Rican relative is T. brenesii Standl., which agrees in pedicellate flowers and an only sparsely puberulous corolla but has much longer (4 mm) corolla lobes and a long-petioled leaf with 15 secondary nerves on each side. LITERATURE CITED NowiCKE, J. W. 1969. Boraginaceae. In Robert E. Woodson, Jr. & Robert W. Schery, Flora of Panama. Ann. Missouri Bot. Gard. 56: 33-69. & . SKVARLA. 1974, palynological investigation of the genus Tournefortia ( Boxasinaceae ). Amer. J. Bot. 61: 1021-1036. NOTES NEW COMBINATIONS IN EPILOBIUM (ONAGRACEAE) In the course of preparation of a revision of the North American species of Epilobium, we have found the new combinations proposed in this paper to be desirable. They will be fully discussed and justified in our subsequent publica- tions, but are offered at this time, with minimal synonymy, in order to make the names available. Epilobium ciliatum Raf., Med. Repos. II. 5: 361. 1808. Epilobium ciliatum subsp. ciliatum. E. adenocaulon Hausskn., Oesterr. Bot. Z. 29: 119. 1879. E. 5 Hausskn. var. macounii Trel., Annual Rep. 1 B. Gard. 2: 103. 1891. achycarpum sensu Munz, Aliso 4: 489. 1960; N. Amer. Fl, ser. 2, 5: 218. 1965, non Presl 1831 Epilobium ciliatum subsp. glandulosum (Lehm.) Hoch & Raven, comb. nov. Based on E. glandulosum Lehm., Stirp. Pug. 2: 14. 1830 E. boreale Hausskn., Monogr. Epil. 279. 1884. E. exaltatum sensu auct. mult.; non Drew, Bull. Torrey Bot. Club 16: 151. 1889. Epilobium ciliatum subsp. watsonii (Barbey) Hoch & Raven, comb. nov. Based on E. watsonii Barbey, in Brew. & S. Wats., Bot. Calif. 1: 219. 1876. Epilobium hornemannii Reichenb., Icon. Crit. 2: 73, fig. 313. 1824. Epilobium hornemannii subsp. hornemannii. Epilobium hornemannii subsp. behringianum (Hausskn.) Hoch & Rave comb. nov. Based on E. behringianum Hausskn., Monogr. Epil. 277. 1884. Support from the U. S. National Science Foundation to Peter Raven is gratefully acknowl- dged. —Peter C. Hoch and Peter H. Raven, Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. CHROMOSOME NUMBER IN PILLANSIA (IRIDACEAE) The chromosome number in the South African monotypic genus Pillansia was previously reported (Goldblatt, 1971) as 2n — 44. New material obtained subsequently proved without doubt to be 2n — 40 (Fig. 1). The earlier record was obtained from paraffin sections of root tips and from anther squashes where 22 bivalents were noted. Plants in the earlier study were all from Rooi Els, Bettys Bay, Cape Prov., South Africa, [Goldblatt 471 (BOL)], and those used in the present work were from Arieskraal, Cape Prov., South Africa, [Powrie s.n. 1977] NOTES 137 Ficuns J. Metaphase chromosomes of Pillansia templemanii stained with lacto-propionic orcein; x 1,200. (MO)] some distance away. A squash technique was used with the new ma- terial, and root tips were treated as described elsewhere (Goldblatt, 1976). Pillansia is a monotypic genus of Iridaceae belonging to the exclusively. Old World and predominantly African. subfamily Ixioideae. The only species P. templemanii ( Baker) L.Bolus is rare and occurs in a very localized area of the southwestern Cape Province of South Africa. Although it is undoubtedly à mem- ber of the Ixioideac, it is peculiar in this subfamily in several respects and hence of particular interest. Most remarkable is its paniculate inflorescence, quite un- like the spike, typical of the subfamily. The branched panicle is believed to be an ancestral condition from which the spike was derived, which suggests that Pillansia is a primitive Ixioid. A second peculiarity of Pillansia is that the corms are persistent, lasting several seasons instead of being annual as is usual in the Ixioideae. Lewis (1954) suggested that this condition in Pillansia was possibly transitional in the evolution of the corm from a rhizome. Thus Pillansia, primi- tive in both its inflorescence and rootstock, is probably a significant evolutionary link between the Ixoideae and less specialized subfamilies of Iridaceae, or their ancestors. The chromosome number of 2n = 20 reported here confirms the polyploid con- dition in the genus. The closest allies of Pillansia—Watsonia, Thereianthus, and Micranthus which together comprise subtribe Watsoniinae (Goldblatt 1971 )— are in contrast basically diploid, with x = 10 in Thereianthus and Micranthus and x= 9 in Watsonia. As it is unlikely that Pillansia is heteroploid, the earlier report of n= 22 appears erroneous. Unfortunately, no more material is available at present to investigate the situation more fully. 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 LITERATURE CITED GOoLDBL ATT, P. 1971. Cytological and morphological studies in the southern African Iridaceae. S. African Bot. 37: 317—460. 976. Chromosome number and its significance in Batis maritima (Bataceae). J. 1 Arbor. 57: 526-530. Lewis, G. J. 1954. Nise on of the morphology, phylogeny and taxonomy of the South African Iridaceae. Ann. S. African Mus. 40: 15-113. —Peter Goldblatt, B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. A NEW JACARANDA (BIGNONIACEAE) FROM ECUADOR AND PERU Jacaranda sparrei A. Gentry, sp. nov. Arbor. Folia pinnatim bicomposita, plerumque 13- gri pinnis 13-21-foliolatis, foliolis m longis, 0.5-1 cm latis, apiculatis. Flor calyce fere patelliformi, 5-dentato, coro tile n) supra basim angustatam arcuatam, Hes a was S staminodio exserto, antheris 1-thecatis, ovario puberulo. Fructus ignotus. Tree; branchlets subtetragonal, very minutely puberulous, with whitish lenti- cels. Leaves pinnately bicompound, usually with 13 pinnae, each pinna with a slightly winged rachis and 13-21 sessile, asymmetrically oblong leaflets, thesc 1-2 cm long and 0.5-1 em wide, apiculate, glabrescent above, barbate at least along the base of midvein below. Inflorescence an open terminal panicle, puberu- lous. Flowers with the calyx almost patelliform, shallowly 5-dentate, ca. 2 mm long and 5 mm wide, puberulous; corolla purplish blue, tubular-campanulate above a narrow neck which is conspicuously curved and enlarged toward the base, 2.5-3 cm long, 1.1-1.3 em wide at the mouth, the lobes small, less than 5 mm long, the whole tube puberulous outside, glabrous inside except at the stamen insertion; stamens didynamous, the anthers 1-thecate, the second theca reduced to a minute appendage, each theca 3-4 mm long, the staminode 2.5-3 cm long, subexserted, the middle third and apex glandular pubescent; ovary flattened- ovate, 2 mm long, 2 mm wide, densely puberulous. Fruit not seen. Tyre: ECUADOR. LOJA: Between Panamerican Highway and Zumbi on road to Machala, km. 69, dry quebrada vegetation, 2100 m, 23 Sep. 1967, Sparre 18862 (MO, holotype). Additional collection examined: Peru. piura: Ayabaca, Oct. 1868, Raymondi 1252 (USM) This species is exactly intermediate between J. acutifolia H. & B. and J. mimosi- folia D. Don on the one hand and the J. caucana complex on the other. It has the relatively large leaflets and pubescent ovary of J. caucana Pittier but the pubescent corolla tube of J. mimosifolia. The curvature and enlarged base of the corolla are more pronounced than in J. acutifolia but less so than in J. caucana. Neither of these species has such reduced corolla lobes nor notably exserted staminodes as 1977] NOTES 139 J. sparrei. Jacaranda sparrei is also intermediate geographically: J. acutifolia occurs in the dry inter-Andean valleys of Peru, while J. caucana occurs from the inter-Andean Cauca and Magdalena valleys of Colombia north to Costa Rica. Supported by NSF Grant DEB75-20325 AOL. —Alwyn H. Gentry, Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. PHYLLARTHRON BILABIATUM: A NEW SPECIES OF BIGNONIACEAE FROM MADAGASCAR Phyllarthron bilabiatum A. Gentry, sp. nov.—Fic. 1. A P. madagascariense foliis angustis nervatura indistincta, a P. humblotiano calvce 5-costato, et ab ambabus foliis verticillatis et calyce bilabiato differt. Large tree to 25 m tall and 0.7 m d.b.h., the trunk convoluted with deep verti- cal fissures, the branchlets subterete to subtriangular, glabrous. Leaves verticil- late in 3's, of 2 superposed articles; petiole ca. 1 cm long; basal article very nar- rowly oblanceolate-oblong, cuneate to the base, rounded at the apex, 3.5-7 cm long, 1.5-2.2 em wide, the second article very narrowly elliptic or elliptic-oblong, rounded at the base, obtuse to subacute or emarginate at the apex, 2-7 cm long, 1-2.6 cm wide; drying olive to gray above, brownish beneath, glabrous, coriaceous, the margins strongly revolute, the secondary nerves very obscure, hardly or not at all visible. Infloresence a short terminal panicle, the lateral branches op- posite, each with 1 or 3 flowers; bracts and bracteoles minute, deciduous. Calyx campanulate, strongly bilabiate, 12-13 mm long, 7-9 mm wide, split over % its length (ca. 5 mm), with 5 conspicuous longitudinal ridges, these terminating in minute denticulations, glandular and drying with a varnished surface, otherwise glabrous. Corolla (single mature corolla seen) magenta with the top of the throat darker, the floor of the throat white with yellow ridges, tubular-infundibuli- form, 4.6 cm long, ca. 1.5 em wide at the mouth of the tube, the tube 2.6 cm long, the lobes 1.2-1.5 cm long, puberulous outside and on the lobes and floor of the tube inside, the lobes also glandular-lepidote. Stamens included, the anther thecae divaricate; pistil and disc not examined. Fruit unknown. Type: MADAGASCAR. DIEGO-SUAREZ: Tsaratanana Massif, trail up S ridge of Maramokotro, Andohanisambirano, 2,000-2,500 m, montane cloud forest, 9 May 1974, Gentry 11612 [MO, holotype; P, TAN, Service Forestióre (Madagascar), isotypes]. Phyllarthron bilabiatum is most closely related to P. madagascariense (Boj.) K. Schum. and P. humblotianum Perrier. Its strongly 5-ribbed calyx suggests the former. Its narrow leaves with indistinct venation and revolute mar gins suggest the latter. Neither of these species has whorled leaves. The leaves of P. bilabia- tum are conspicuously decurrent so that its branchlets appear almost triangular ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 140 Ficure I. Habit of Phyllarthron bilabiatum A. Gentry (X %40). [Gentry 11612 (MO).] in cross-section. The most noteworthy floral character of this species is a strongly bilabiate calyx which is matched in the genus only by the very different P. megap- terum Perrier. Perrier de la Bathie (1938a, 1938b) noted the variability of juvenile leaves of this genus and excluded them from consideration in his key and species descrip- tions. I have followed suit in using only the mature foliage in the description of the new species. However, a sterile collection from the type locality [Gentry 11618 (MO) described as a sterile treelet 2 m tall] certainly represents a juvenile form of P. bilabiatum. The leaves of this collection are larger and thinner than 1977] NOTES 141 mature leaves and have secondary venation approaching that of P. madagas- cariense, but their whorled placement agrees with P. bilabiatum. The stem of this plant is distinctively triangular from the strongly decurrent leaves, a charac- ter which appears to distinguish juvenile plants of P. bilabiatum from juvenile forms of any other species of the genus. LITERATURE CITED PERRIER DE LA Bartnik, H. 195358. ^i D de la region Malgache. Ann. Inst. Bot.-Géol. Colon. Marseille, sér. : 110 1-18. 938b. Bignoniacées, In n bed [ye Flore de Madagascar, 178° Famille. 9] pp. —Alwyn H. Gentry, Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. TAXONOMIC NOTES AND NEW COMBINATIONS IN LEUCOPHYSALIS (SOLANACEAE) During the course of a revisionary study of Chamaesaracha (Averett, 1973), several species were encountered which, at one time or another, had been assigned to but are clearly not a part of Chamaesaracha. Most of these species were relegated, either by me or previous workers, to Leucophysalis or the Asian genus Physaliastrum. In dealing with the misplaced species, the close relation- ship of Physaliastrum to the North American genus Leucophysalis became ap- parent, but since the species were removed from Chamaesaracha, the question of the two being congeneric was postponed (Averett, 1971). The data now at hand indicate that the species of Physaliastrum are clearly related to and are best treated as Leucophysalis. The latter treatment necessitates scveral nomenclatural changes. Since my revision of the genus will not appear for several months, it seems advisable to make the following new combinations at this time: Leucophysalis kweichouense (Kuang & Lu) Averett, comb. nov. Mari caet y kweichouense Kuang & Lu, Acta Phytotax. Sin. 10: 351. 1965. rype: China, ‘eichou Province, Keili, Maopin, 750 m, Chang Yongtien 1396 (SH, holotype, not seen). Leucophysalis sinicum (Kuang & Lu) Averett, comb. nov. a sinicum Kuang & Lu, Acta Phytotax. Sin. 10: 352. 1965. ‘rype: China, Shansi vince, Wei-ying Hsia 4321 (SH, holotype, not seen). — Leucophysalis yunnanense (Kuang & Lu) Averctt, comb. nov. R yunnanense 17 05 & Lu, Acta Phytotax. 10: 348. 1965. Type: China, Yunnan ince, Sunning, 1800 m, T. T. Yu 16767 (SH, holotype, not seen). Leucophysalis japonica (Fr. & Say.) Averett, comb. nov. (—— japonica Fr. & Sav., Enum. Pl. Jap. 2: 454. 1879. TYPE: Japan, Ito Keiske, anaka, Savatier 2166 phe seen — 142 ANNALS OF THE MISSOURI BOTANICAL GARDEN Vor. 64 C. echinata Yatabe, Bot. Mag. (Tokyo) 5: 355. 1891. Type: Not designated. Physaliastrum echinatum sd Makino, Bot. Mag. (Tokyo) 28: 20. 1914. P. jap m (Fr. & Sav.) Honda, Bot. Mag. (Tokyo) 45: 139. 1931. P. AMD (Fr. & Sav.) E. Acta Phytotax. Geobot. 6: 19. 1937. Leucophysalis savatieri (Makino) Averett, comb. nov. on savatieri 11 15 Illustr. ee Jap. I. (11): 1. Oct. 9, 1891. rEecrorvPk: Japan, Nikko. Oct Makin G. Nau) Yatabe 9 5 oC "folky ah. "315. Oct. 10, 1891 Phusaliastrum savatieri (Makino) Makino, Bot. Mag. 17 8 28: 22. 1914. Leucophysalis kimurai (Makino) Averett, comb. nov. V iy bere J. Tap. Bot. 32-37. m TYPE: Japan, Musashi Province, Mt. 926, K. Kimura s.n. (not seen P: Miren: ud 7 Ohwi, Fl. Jap. 1026. 1956. Nine species in total, two North American and seven Asian, are considered to compose the genus Leucophysalis. LITERATURE CITED AVERE f. 3 New combinations a ho 5 (Solanaceae) oe comments rearing ie taxonomy of Leucophysalis. uri Bot. Gard. 57: 380-382. — O 1973. Binaries study of ape ats (ona. 8 75: 325-365. —John E. Averett, Department of Biology, University of Missouri-St. Louis, St. Louis, Missouri 63121. CHROMOSOME NUMBERS OF PHANEROGAMS. 7' Counts by Charles Albert Huckins, Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. ROSACEAE Malus baccata (L.) Borkhausen. 2n = 34. U.S.A. WASHINGTON, D.C.: Cul tivated, U.S. National Arboretum accession number 2059 (PI 107683), Pal 66042101 (BH Malus eee var. himalaica (Maximowicz) Schneider. 2n = 34. MASSACHUSETTS: ultivated, Arnold Arboretum accession number 0 Huckins 69050813 (BH). Malus florentina (Zuccagni) Schneider. 2n = 34. U.S.A. WASHINGTON, D.C Cultivated, U.S. National Arboretum accession number 3361, Huckins 67081802 Malus fusca ( Rafinesque) Schneider. 2n = 34. U.S.A. NEW york: Cultivated, Durand-Eastman Park accession number 77, Huckins 69052703 (BH). Malus rockii Rehder. 2n = 51. U.S.A. MassacHusETIS: Cultivated, Arnold Arboretum accession number A 83-84, Huckins 70051902 (BH). 1 The previous number in this series appeared in Ann. Missouri Bot. Gard. 62: 513. 1975. 19771 NOTES 143 Malus sikkimensis (Wenzig) Koehne ex Schneider. 2n = 51. U.S.A. NEW york: Cultivated, Durand-Eastman Park accession number 841, Huckins 69052105 (BH). Malus tschonoskii ( Maximowicz) Schneider. 2n = 34. U.S.A. NEW york: Cul- tivated, Cornell University Horticulture Department accession number URI- 12, Huckins 68041701 (BH). Malus yunnanensis (Franchet) Schneider. 2n = 34. U.S.A. NEW YORK: Cul- tivated, Cornell University Horticulture Department accession number URIA, Huckins 69050601 (BH). The previous issue of the ANNALS or THE MISSOURI BOTANICAL GARDEN, Vol. 63, No. 4, pp. 657-896, was published on 14 June 1977. pu 3 E. A. Bu rt ee | "Publications c of the ` MISSOURI BOTA CAL GARDEN 5 The Monts. OF THE . | RUN 98 contains con- ee s tributions mainly in plant systematics. The ANNALS appears four times a year, and four numbers constitute a volume. Many back issues are — The ee seated are aod net; there j is no disco count to ¿ Dt 2 rimarily systematics, Dut early volumes contain i numerous papers on topics ranging from n - _ Azotobacter to pod and stem blight of soybean. Volumes 1-15 con- a taina series of BS papas 3n Noh Anidos Th by B. M. publis ed š | volume8, — pe e first 55 eM aus (16141088) of the PU are now E dei; The INDEX catalogues the 769 titles contributed by 326 authors i in these volumes. The IN NDEX i is s soft bound and costs ess 00. | BOTANICAL GARDEN 1977 NUMBER 2 TLLA — ca I Arg ELE. tUm = — ae es VVV "FER. og ES à š | CONTENTS GARDEN LIBRAR CC The Twenty-third Systematics Symposium John E. Averett 145 Perspectives in Plant Serotaxonomy David E. Fairbrothers 147 Electrophoretic Evidence and Plant Systematics L. D. Gottlieb 161 The Applications of Molecular Evolution to Systematics; Rates, — tion, and the Role of Natural Selection Mary-Claire King „ Chemosystematics Analysis of Populational Differentiation and V js d of Ancestral and Recent Populations of Juniperus ashei Robert P. NEN oa o ñß Re ñ cu uei m iM e agre 184 The Order Centrospermae Tom J. Mabry / 0 Defensive Ecology of the Cruciferae Paul Feen 221 Chemosystematics and its Effect upon the Traditionalist B. L. Turner .. 235 (Contents continued on back cover) VOLUME ó4 1977 NUMBER 2 OF THE MISSOURI BOTANICAL GARDEN The ANNALS contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden. Papers originating outside the Garden will also be accepted. Authors should write the i editor for information a ss for ‘Publishing à in the ANNALS. | Eprronta AL . Commrrree | W. G. Dass Editor— Flora E Panama aes Missouri Botanical Garden ; i Jonn D. DwYER oe Missouri moe Garden & St. a eran Gol b BLATT : ` (un Botanical Garden — Published four times a year bs the Missouri Botanical Carden Press, St. Louis, Missouri 63110. For Subscription information contact the Business Office of the Annals, . Box 368, 1041 New Hampshire, Lawrence, Kansas 66044. Subscription price is $40 p per volume U.S., Canada, = Mexico, 845 all other countries. Four issues per volum Second me postage paid a at Lawrence, Kansas 66044 ANNALS OF THE MISSOURI BOTANICAL GARDEN VOLUME 64 1977 NUMBER 2 CHEMOSYSTEMATICS: THE TWENTY-THIRD SYSTEMATICS SYMPOSIUM JOIN E. AVERETT! The following seven papers were presented at the Missouri Botanical Gar- den’s Twenty-third Annual Systematics Symposium held 15-16 October 1976. The symposium, sponsored in part by the National Science Foundation, was at- tended by approximately 300 scientists. This year’s topic was chemosystematics. The studies of R. E. Alston and B. L. Turner in the early 1960s which utilized flavonoid patterns in elucidating complex hybridization and their publication of Biochemical Systematics in 1963 were instrumental in bringing chemistry to the systematic community. In 1962 the first international conference on biochemical systematics was held, and the proceedings were published the following year in Chemical Plant Taxonomy (Swain, 1963). Taxonomic Biochemistry and Serology, also proceedings of an international conference held in 1962, ap- peared in 1964 (Leone, 1964). Through the 1960s numerous papers dealing with natural products appeared in the literature and by 1970 the field of bio- chemical systematics or chemotaxonomy was well established. In 1972 the International Union of Pure and Applied Chemistry held a sym- posium in Strasbourg on “Chemistry in Evolution and Systematics” (Swain, 1973). In the following year the topic of the 25th Nobel Symposium was “Chemistry in Botanical Classification” (Bendz & Santesson, 1974). The latter was an attempt to bring chemists and taxonomists together, in what V. H. Hey- wood has referred to as the unlikely marriage of an exact science with one less restricted that ventures into every other discipline. Early studies utilized compounds, and except for serology, largely secondary compounds, in resolving hybridization complexes or as “finger prints” for charac- terizing species or, occasionally, higher taxa. However, concomitant with the development of the field, techniques and instrumentation necessary for the iso- lation and structural characterization of the compounds were refined. Further, ! Department of Biology, University of Missouri-St. Louis, St. Louis, Missouri 63121. Moderator of the Twenty-third Annual Systematics Symposium. ANN. Missouni Bor. Garp. 64: 145-146. 146 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 information on biosynthetic pathways and genetics of the secondary compounds accumulated. Now it is relatively easy to assay the complex constitution of a plant. Structural and biosynthetic data greatly enhance the utility of the com- pounds in phylogenetic interpretations. The extent to which this is true, as well as the sophistication that has developed within the field, is well illustrated in the following papers. Following the format of the symposium, the first three papers discuss the utility of macromolecules and next three papers discuss micromolecules. The final paper on the effect of chemistry upon traditional taxonomists was presented as an evening talk by B. L. Turner. LITERATURE CITED ALSTON, R. E. . L. 1 5 1963. Biochemical Systematics. Prentice Hall, Inc., En- p us. New Jers BENpz, G. ANTESSON editors). 1974. Chemistry in p ero Classification. Proc. 25th Nobel Symposium, 1973, Sweden. Academic Press, LEONE, C. A. (editor). 1964. Taxonomic Biochemistry and 8 Ronald Press, New York. SwAIN, T. 1 editor). 1963. Chemical Plant Taxonomy. Academic Press, New York. a 1973. Chemistry in Evolution and Systematics. Butterworth & Co., Ltd., London PERSPECTIVES IN PLANT SEROTAXONOMY! Davin E. FAIRBROTHERS” ABSTRACT The capacity to view recent data in proper relation to other information, or the abili ty to correctly judge the significance of facts and ead requires a knowledge of both the past mistakes and the forward strides within a disc cipline. This paper is intended to help the reader formulate perspectives concerning 65 years of plant serotaxonomic research. The discovery that the immune reaction was only relatively specific and that the degree of cross- reactivity was essentially proportional to the degree of PE po 'en organisms had important bea for comparative systematic serology. It the specific reactions, between determinants anc fo^ denis which Dione a WM cox of protein simi- larities. The comparison of protein mixtures, ra the than purified single proteins, has dominated taxonomic research because such an approach provides ser ological overall similarity, and thus a multicharacter comparison. The “antisystematic” reactions have recently been shown to result from variation in the systematic ! ranges of detern + ad and the absorption (presaturation) technique for removing common determinants increases the accuracy of serological placements. ae following items were evaluated: antigenic preparations, ad- juvants, injection procedures, single versus mixed Nene extractions. kind of plant tissue extracted, and the interference of secondary 5 Cornus canadensis and C. suecica were found to be serologically very similar. The tested species of the genus Cornus were divided into three distinct serological groupings. The serological data support the separation of the Cornaceae and Nyssaceae; and the inclusion of Camptotheca anc Nyssa in the Nys- soideae, and Davidia in the Davidioideae, both of the family Nyssaceae. Nyssa biflora and N. sylvatica were serologically very similar; N. ogeche and N. aquatica were serologically distinct from each other and from N. biflora and N. sylvatica. Nyssa ogeche was the most dis- tinct species of the genus. Corokia cotoneaster had very little serological similarity with any of the tested species of the Cornales. To have the capacity to view recent data in proper relation to other infor- mation, or the ability to correctly judge the significance of facts and ideas, re- quires a knowledge of past mistakes and the forward strides within a discipline. It also requires a degree of knowledge of the individual components as well as the total products resulting from the various component combinations. The author hopes this manuscript will provide the reader with the necessary infor- mation and literature citations which will allow the formulation of perspectives concerning 65 years of plant serotaxonomic research. The “present age” of chemosystematics or chemotaxonomic publications com- menced to appear in the early 19508. The oldest of the “present age” approaches is serotaxonomy and the newest is amino acid sequencing (Cronquist, 1976). The discovery of serological reactions in Austria via the occurrence of precipi- tin reactions took place 80 years ago (Kraus, 1897). This discovery provided a new technique which was soon used to aid in the investigation of systematic problems in animals. Within two years after the discovery of the precipitin re- dedicate this publication to the memory of my friend and fellow F Dr. P es who died in his 55th year on October 22, 1976 in Czechoslov The research was supported by NSF Grants GB- 13202 and BMS 75-17805. * Department of Botany, Rutgers University, New Brunswick, New Jersey 08903. ANN. Missouni Bor. Garp. 64: 147-160. 148 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 action the technique was applied to comparative problems by the Frenchman Bordet (1899). This was followed by a series of extensive comparative studies which were conducted with various animals by the Englishman Nuttall (1901, 1904). Thus biologists have known for 75 years that organisms may share anti- genic material (substances capable of inducing the formation of antibodies and able to react with the antibodies); and that when they share the same anti- genic material in different proportions it is assumed that the organisms are re- lated. At first it was believed that the immune reaction was absolutely specific, that is, that an antiserum would react only with the antigen that stimulated its production. However, Bordet (1899) in conducting research with birds reported that the reaction was only relatively specific and that the degree of cross-reactivity was essentially proportional to the degree of relationships between organisms. It was this early discovery which had important implications for systematics and started the pioneer investigations in the discipline of comparative animal sys- tematic serology. large number of animal systematic serological publications have been re- ported in the bibliography prepared by Leone (1968) and the book edited by Wright (1974). The pioneering work in the United States essentially began with Boyden (1926), and he has continuously contributed to the field of animal systematic serology for 50 years (Boyden, 1973; Wright, 1974). Approximately 550 plant taxa (cultivars through orders) have been included in approximately 160 systematic serological publications since 1950 (Fairbrothers, 1969a; Fair- brothers et al., 1975). Serology is concerned with the interactions of antigens and antibodies and/ or antibodylike substances, the lectins. The term "serology" is often used synony- mously with immunology. However, some biologists prefer "serology" because “immunology” has an implication that immunity is concerned in all reactions be- tween antigens and antibodies ( Boyden, 1948). PRINCIPLES OF SEROLOGY A consideration of a few basic facts related to the biology of the immune response is a valuable aid to understanding the methods and interpretations of systematic serological research. e term “antigenic” is relative since the response is frequently a property of the route of injection, method of preparation of the antigenic material, and the individual experimental animal used. Thus it is important that details about such items and procedures should always be included as a portion of the methods and materials section of publications. Experimental data have demonstrated the following: (1) There is a certain minimal molecular weight below which substances are not in themselves antigenic. Some of the lower molecular weight substances can become antigenic by mixing them with other substances (ad- juvants). (2) Size alone is not enough to guarantee that a molecule will be antigenic. Immunochemists indicate that the specific action is in part due to the rigidity of certain chemical structures (determinants) which are difficult to distort or alter. (3) Usually a molecule must be foreign to an organism to be immunogenic. (4) Substances must be soluble or be able to be broken down 1977] FAIRBROTHERS—PLANT SEROTAXONOMY 149 into soluble antigenic components before being capable of inducing antibody formation. (5) Too much antigenic material may cause immunological paralysis (i.e., cannot be immunized). There appears to be a balance between stimula- tion and paralysis for each antigenic material. (6) Many proteins have been found to be immunogenic, and the best known immunogens are the proteins with molecular weights of 40,000 or more (Abramoff & La Via, 1970). Of the several kinds of serological reactions, the precipitin reaction has been used most frequently in plant comparative serological investigations. It is a rela- tively simple reaction capable of being applied to the comparison of the soluble protein antigenic material extracted from all kinds of plants. Microcomplement fixation has proved valuable in animal systematics (Champion, et al, 1974), but has had practically no application in comparable plant research. Serological research can be conducted employing quantitative precipitation, precipitin tech- niques in solutions, or by various qualitative precipitation techniques in gels. The serological characteristics of proteins are linked with the primary structure of the molecule. The reaction is concerned with points on the molecule ( determi- nants) which are capable of initiating the production of immuno-globulins only in certain cells of animals (not plants). These immuno-globulins possess proper- ties accounting for the bonding to the respective protein reaction position. Thus the serological characteristics of the protein are found in the determinants, which are restricted to certain positions of the molecules. fairly accurate estimation of the size of determinants has been obtained from protein fractionation experiments designed to detect the smallest molecule fraction still capable of an immunological response. Arnon & Sela (1969) and others have demonstrated that the active antigenic regions of proteins are com- posed of 10-20 amino acids. Systematists and taxonomists are interested in the comparison of antigenic determinants from various taxa. It is the specific reactions between determinants (antigens) and antideterminants (antibodies) which are valuable because they provide a means for the measurement of protein similarities. METHODOLOGY When deciding the type of antigenic preparation, the process of denatura- tion, which means structural changes with concomitant loss of biological proper- ties, must be taken into account. Protein antigens are not equal in susceptibility to denaturation. However, all such changes result in some loss of original specific- ity. The use or nonuse of adjuvants ( Freund's, in our experiments) to increase the level of an immune response (immuno-enhancement) is discussed in systematic serological research. The purpose of an adjuvant is to heighten and prolong the immune response; and the value of this additional material must be judged for sach antigenic material. This means its use or nonuse should be decided after experimentation. The effect of injection procedures on the systematic reaction range has been tested by various experiments. One of the very early reports indicating that longer immunization periods extend the reaction range was conducted with a Zea mays 150 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 antiserum ( Magnus, 1908). Lake et al. (1914) using purified seed protein ma- terial in contrast to crude seed extracts as used by Magnus also found that a longer injection period extended the reaction range of the antisera. Several recent publications, with both plant and animal antigens in the form of purified or mixed reagents, indicate that the systematic reaction ranges of antisera are extended by injections continued until a maximum reaction is reached (Boyden, 1971, 1973). Such experiments indicate also that long continued immunization is likely to induce the formation of greater proportions of cross-reacting anti- bodies, which will reduce the discriminating capacity of the antiserum. Antisera derived from several and long injection periods may reach a higher level of "fidelity" to mixed antigens, and reveal information of value in systematic re- search (Moritz, 1964). Leone (1952) demonstrated that longer immunization periods tend to cause a lower discriminating capacity. Therefore, the number of injection series used should be stated so the reader can make proper comparisons. The use of a combination of “short” and “long” injection series may produce the largest amount of data for comparative sero- logical investigations. This should not be considered experimental “manipula- tion" because it is merely using serological techniques to the fullest advantage based upon our present knowledge of the immune response. There are two main approaches to serological research: (1) comparison of single proteins, or (2) comparison of protein mixtures. The second approach has dominated taxonomic research (Moritz, 1964; Fairbrothers, 1968; Jensen, 1974a). An example of the comparison of a single protein is best illustrated by the re- search with phaseolin obtained from Phaseolus vulgaris, and the ribulose-1,5 biphosphate carboxylase ("fraction I") found in green plants. Data obtained from such research indicates that the systematic ranges of determinants vary. Some determinants are found throughout the plant kingdom, while others have a very restricted distribution. In general, the low taxonomic yield from single protein investigations has not justified the large preparation effort required (Jensen, 1974a). Serological comparison of protein mixtures which were ex- tracted from seeds, pollen, spores, tubers, or leaves is most common. The re- sults provide a serological overall similarity and thus a multicharacter compari- son. Researchers working with plant protein extracts also find tannins, saponins, alkaloids, lipids, and/or polysaccharides, which may have to be inactivated, re- duced, or eliminated by diverse extraction procedures. When using a pure protein, only a very limited number of determinants can be compared. In contrast, when using mixtures of proteins, data from many dif- ferent determinants are tested, and thus the chances of being misled in terms of serological correspondence are lessened. "ANTISYSTEMATIC REACTIONS > << A phenomenon which has been designated “antisystematic,” “asystematic,” or “unexpected” cross-reactions has only very recently been placed in proper per- spective ( Moritz & Rohn, 1956; Frohne et al., 1961; Moritz, 1964). Jensen (1974a, 1974b) indicated that these terms are no longer used by the above authors be- cause past usage assumed serological convergence, which has been shown not to 1977] FAIRBROTHERS-—PLANT SEROTAXONOMY 151 be the causative agent for such responses. Moritz (1964) included several re- ports which illustrated his designated "antisystematic" reaction. He also indi- cated why such reactions were specific serological reactions, and not some kind of non-specific effect, since they disappeared when presaturation experiments were conducte he practice of some researchers during the last several years of designating cross-reactions with a wide systematic range as "antisystematic" should be dis- continued. Such wide-range reactions are the result of certain determinants be- ing widely distributed in the plant kingdom. In other words, they should be con- sidered as reactions of determinants that are widespread and remain relatively constant. One such determinant has been demonstrated by the serological re- search with Fraction I Protein (ribulose-1,5 biphosphate carboxylase) extracted from the tissue of green plants. The serological partial identity detected between wide-ranging taxonomic groups has been shown to be the result of parts of the protein structure unaltered in the course of long periods of evolution (Sugiyama et al., 1969; Jensen, 1974a, 1974b). Thus we now know that the systematic ranges of determinants do vary, and sometimes protein molecules carry several de- terminants which reveal partial serological identities. The understanding of the above reactions is important because it has shown the value of the presaturation (absorption) technique for removing common determinants and leaving only those systems specific for each taxon compared, thus providing both a more accurate serological placement and measure of the relative similarity. The use of various techniques to remove nonspecific reactants which react with normal rabbit serum (NRS) has become standard practice in present-day plant serological research. We now know such responses often come from sero- logical reactions resulting from the presence of lectins. Lectins can be removed by hemagglutination techniques and thus be prevented from interfering with normal serological reactions (Lee & Fairbrothers, 1972). us in recent years experiments have provided answers to some of the for- mer perplexing problems associated with plant systematic serological research. This has been very valuable and allowed the continued development of such research. It also assures that a larger spectrum of species can be compared by using the techniques which are now available, and our percentage of accuracy in terms of serological placements, is continuing to increase. History OF PLANT SEROLOGY As with animals, serological techniques were used in plant systematics and taxonomy soon after discovery. Mez of the Botanical Institute, University of Königsberg, Germany, conducted such research with his students and colleagues from 1911-1936 (Mez & Gohlke, 1914; Mez, 1922). The "Kónigsberger Sero- diagnostik Stammbaum” (phylogenetic tree) was the climax of years of research (Mez & Ziegenspeck, 1926). This group was known as the Kónigsberg Sero- logical School, in contrast to the Berlin Serological School which was headed by Professors Gilg and Schiirhoff, who conducted research during the 1920 (Gilg & Schiirhoff, 1926). These two groups of researchers (schools) conducted 152 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 literary feud which seriously jeopardized the credibility of plant systematic serological research for essentially 25 years. The techniques employed in the early research proved to have several serious flaws. Mez's technique, where he produced an immune serum through the influence of antigen on serum and eliminated the use of living animals as antisera producers, was a serological "disaster." The vast amount of data reported using this procedure was not valid and has been disregarded. It was Moritz (1928, 1964) and his colleagues at the University of Kiel, Germany, who, working from 1928 until the present, revealed the value of sero- logical research for plant systematics. Jensen, formerly from Kiel, has recently organized another serological laboratory at the University of Cologne. Frohne continues the systematic serological research at Kiel. In the United States Chester published plant serological papers in the 1920's and 1930's and also prepared a comprehensive critique about plant systematic serology (Chester, 1937). In 1947 Johnson (1951), with his students, started the present United States trend toward plant systematic serological research. It was he who introduced me to the techniques in 1957 after I had joined the faculty of Rutgers University. In 1953 Urano (1955) started phytoserological investigations in Japan, and S. Sakaguchi and S. Arai have continued this research (Fairbrothers, 19694). The Solanum serological research in Birmingham, England was started in 1955 (Gell et al., 1956). J. Hawkes (Birmingham) and his students (Lester and P. M. Smith) have continued serological research to the present time. In Prague, Czechoslovakia in the late 1950's the husband and wife team, Kloz and Klozova, started and continued serological investigations of Vicia faba and other legumes (Kloz et al, 1960). The year 1963 saw the start of another plant serological research center headed by Vaughan in London, and he has continued his multi- disciplinary Brassica investigations to the present time (Vaughan & Waite, 1965; Vaughan et al, 1976). Cristofolini (1968) has published several reports from his botanical laboratory in Italy since beginning his plant serological research in 1966. The most recently organized plant systematic serological laboratory is under the leadership of Drs. Morozova, Chupov and Kutjavina at the Komarov Botanical Institute, Leningrad, USSR (Fairbrothers, 1975). INTERPRETING THE RESULTS Research has demonstrated that extracts of seeds, pollen, leaves, tubers, and spores of vascular plants can be used, if the required extraction procedures are followed (Fairbrothers, 1969b, 1975; Fairbrothers et al., 1975). However, most systematic serological research has included seed material, due to the relatively high concentration of proteins, relative ease of collecting, and relative ease of assuring comparable developmental stages. We are presently pursuing the fol- lowing two new studies in our chemosystematic laboratory using pollen as the source of protein material: (1) serological investigation of selected amentiferous taxa with Frank Petersen, and (2) a serological investigation of the Corylaceae ( Betulaceae) with Friedrich Brunner. 1977] FAIRBROTHERS PLANT SEROTAXONOMY 153 Phytoserological research has provided provocative and valuable data for use in the classification of flowering plants. The numerous examples cited in the evaluation of the contribution of serological data related to Cronquist’s and Takhtajan’s systems of classification were shown to be significant (Fairbrothers et al., 1975). This publication indicates that such data have contributed in the classification of the following orders, and the placement of families within these orders: Capparales, Caryophyllales, Cornales, Dipsacales, Illiciales, Lamiales, Magnoliales, Nelumbonales, Nymphaeales, Papaverales, Polemoniales, Ranun- culales, Rubiales, Scrophulariales, Typhales, and Umbellales. In addition to these orders, significant contributions have also been published for species, genera, and/or tribes belonging to the following families: Ammiaceae, Ber- beridaceae, Brassicaceae, Caprifoliaceae, Cucurbitaceae, Chenopodiaceae, Cor- naceae, Fabaceae, Lamiaceae, Magnoliaceae, Nelumbonaceae, Nymphaeaceae, Nyssaceae, Papaveraceae, Poaceae, Ranunculaceae, Solanaceae, and Typhaceae. The serologic and disc electrophoretic characterization and comparison of the spore proteins extracted from Osmunda cinnamomea, O. claytoniana, and O. regalis illustrated that fern spores were suitable material for such analyses. Osmunda cinnamomea and O. claytoniana were shown to possess greater protein affinities for each other than either had for O. regalis. Osmunda regalis, in gen- eral, had greater protein affinities for O. claytoniana than it had for O. cin- namomea (Petersen & Fairbrothers, 1971). Stein & Thompson (1975) compared the same three Osmunda species using DNA hybridization techniques and in- dependently indicated the same relationships reported by Petersen & Fairbrothers (1971). Miller (1967), based on anatomical characters of living and fossil speci- mens, indicated that O. claytoniana and O. regalis were more closely related, while Hewitson’s (1963) anatomical and morphological research indicated that O. cinnamomea and O. claytoniana had a closer relationship with each other than either had with O. regalis. The serological investigation of intra- and interfamily relationships of the Cornaceae and Nyssaceae has continued intermittently in our chemosystematics laboratory for 15 years, and various experiments have been conducted as ap- propriate and adequate plant materials became available. In our systematic serological research it has been expedient to conduct several projects simultane- ously because no experiments can be conducted until adequate and appropri- ate materials are available for the extraction of proteins, and until antisera to perform the essential experiments have been raised. Cornus canadensis and C. suecica have been found serologically very similar based on photronreflectometer tests, Ouchterlony plates, and absorped and non- absorbed antisera. These two taxa have also been shown to be the most dis- similar from other taxa placed in the genus Cornus (Fairbrothers, 1966a, 1966b, 1968). When data were evaluated from cytology, morphology, anatomy, geo- graphical distribution, and the putative hybrid (C. unalaschkensis), the close similarity between the two was also detected. I believe all the data indicate that — the two named taxa are subspecies of one circumboreal species which is very distinct from the other species of Cornus. If there is justification for dividing the genus Cornus into distinct genera, then this species (or two species) would [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN “paalaoal [eurrue [equəurriiədxə dove serias uonoefur jo 1əqumu əy} 9jeorpur siequmnu rjosque əy} uo sjduosiedns L q (8961 49961 '*996I “siəqqoriq -nur fpgeI *uosuqof X Sm q)) exe} pepn[our əy} jo əuuos 10j ejep 193jaurojoo[]joruomoud peusmqnd SnOIA91d dum pojen[eAo aq ?jep Aou əsəqn1 FLY} [enuəssə SI J v S EY r 01 01 — OI € E zm Ë 49}SDIU0}JOI 0 001 — r€ — IF — 9 18 == — £c Djpionjoaut d GS 001 001 88 88 56 0G FE cI 18 88 Donupajfis N EE 98 — 001 68 0G LS [4i 9c xus 2u9280 'N ET 66 99 9L 001 001 08 — SI 9c 98 vonpnbD N 68 89 FP LY 0€ 69 88 pE Sc 88 08 DIDUIWNID e —_ 86 — cI — ce OF 5 001 88 = sisuəppup2 :D — — — — — — c9 — — - 0 nppanu 72) — — — == = us 09 = == = 85 vn x — m — yc — c9 — m — 58 ppuio]f `D — SFr 96 86 EE LS 001 001 6f v8 €L DSOUL2ODA `D) 5 m — — — — E — mE — 18 Diafiuojo}s `J — LY LG c€ TE 9S d 06 OS 001 001 tumuotup ‘D :6017H :9II-H FIDCM II- N III M 86-H SCI rC IM «901-H :SITI-H -0TI-H suadnuy DIDIN OAU? pOpajfis"N — DOpajfis N 2429280 ` N D2MgpnbD'N D2DnbDD^N DSOU4299D4 `D USOUL29DA4 'O SISUJDDUDO ^) WNUWOWD "[) | Ulm utQutD 7j) ‘d qeissquy 2001 SE passaidxea st porga *u0rj?al ou ƏY} JO vorn juoo1od juosai1dai siəqumu AL '19PPUOPƏAJu01704d Əy} Bulsn (N) DssfiN pue () Dipiang ‘(09 ) myorop (`D) nul ‘('89) 722130101407) jo səpəds yy suonoeai untrdrooid woy pəurezqo PA I TYL FAIRBROTHERS—PLANT SEROTAXONOMY 155 1977] 'sioqpjoiqure q :f96I “uosuuoíf “spueq JO Joquinu [v}0} = L [d [d 0 I U 0 [A 0 F 8 g I ° I [d S 0 S ee 8 I C 8 I rd € 0 € 8 I G e I G C 6 0 6 C 0 ra I I [4 I I [d I I 6 I I L oN I L N I SOLA ITTY Djnionjoau: ` D911pq]ñs `N ‘spueq 4jQquopr pened 10/pue Ajguopruou = N Pole [eurrue. [e3uourniodxo qovo sorios 7? sioqyoiqie) vxej pəpnpur oy} jo Əuuos 10j ep [eotSo[o1os ‘spueq Xjquapr = I *SAO[[0J se PayeUSIsap uəəq savy suiojged Surpurg 5 uonooefur jo Jaquinu ay} ajeorpur sriaquinu viesgur aug] uo sjduosiodus SUL q '(896I 49961 Peusiqnd snoraaid qjr^ pojen[eAa aq VPP Mau ISIYI jeu] [PUUƏSS3 SI JJ = 'tg961 I I ra ra 0 T T 0 I I 0 I I 0 4228D9u0]00 í 3 — UM r4 I I 55 r DJDAINJOAUL d B F Y. 8 I Ë 0 I I 0 ¢ I I Doijpa]fis "Nr F O € I c ¢ € 0 I I 0 € I I au93850 "N € 0 7 O 5 ¿€ Z 0 I ] 0 8 l I D21]pnbp ^N € I © I 8 1 I I I 0 € I l DIDUIUINID “DD I I T = 0 T I I FP O F e € O sIsuəppupa 7) T € € I I F 0 * € I I € 1 ra DSOWIIDL `D e 8 c I I € I T r4 I l € 0 € utnutotup Z Y L N L N I L N I L N I L N I suog Huy We Sous uu I-A :867H ELN "TENE mora 342230 'N Dongpnbp'N DSOWABIDA `D SISUIPDUDI "FD UTIULOUID )) qeiosquy `suədnue əutu YAN pojeduoo oie LIƏSIJUL UAIG (N) Ds pue (A) piang (CO) nyoro -ojduupo *('D ) snusog 10} (uotsnjjip Ə|qnop ) soje[q Xuo|jiə1uonO uro1j pəuwy qo (spueq) suujsás Bunezrdoordounwun jo Jaquiny rg) 5001 „ C ATV 156 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 best qualify for such a designation, and would have to be given the generic name of Chamaepericlymenum. Serological data have also indicated that within the genus Cornus there are the following three distinct groupings: (1) C. florida, C. kousa, and C. nut- tallii; (2) C. amomum, C. stolonifera, and C. racemosa; and (3) C. canadensis and C. suecica ( Fairbrothers & Johnson, 1964; Fairbrothers, 1966a, 1966b, 1968). These serological groupings correspond to the Cornus subgenera designated by Ferguson (1966), except he placed C. kousa in a subgenus distinct from that containing C. florida and C. nuttallii. Newer serological data from our lab- oratory based upon additional antisera and many more experiments continue to support a tripartite taxonomic disposition of the taxa placed in the genus Cor- nus (Tables 1-2). The use of nonflavonoid glucosides as taxonomic markers in the genus Cornus was reported by Jensen et al. (1975). Their suggested ar- rangement of subgenera based on the presence or absence of iridoids, plus the type of iridoid constituents agree with the reported serological groupings and would correspond to their (A/B), (C), and (F/G/H) designations. The families Cornaceae and Nyssaceae were recognized by Dumortier in 1829. However, this separation into two families was not followed by most taxonomists for over 100 years. Recently Melchior (1964), Cronquist (1968), Thorne (1968, 1976), Takhtajan (1969), and Dahlgren (1975) have recognized two families. In addition, in most recent classifications the genus Davidia has been removed from the Nyssaceae and placed in the Davidiaceae (Melchior, 1964; Cronquist, 1968; Takhtajan, 1969; Dahlgren, 1975). Thorne (1968, 1976) placed Davidia in the subfamily Davidioideae of Nyssaceae following Wagerin (1910). Harms (1898) was the first author to use the two subfamilies Davidi- oideae and Nyssoideae, and he placed them both in the family Cornaceae. The serological data support the separation of the Cornaceae and Nyssaceae, and the grouping of Camptotheca, Davidia, and Nyssa within the Nyssaceae (Tables 1-2). At present I believe the serological data best support the place- ment of Davidia in the Davidioideae, and Camptotheca and Nyssa in the Nyssoideae both of the family Nyssaceae (Fairbrothers & Johnson, 1964; Fair- brothers, 1966a, 1966b, 1968; Tables 1-2). Perdue et al. (1970) reported that tests with Camptotheca acuminata dem- onstrated that crude extracts exhibited significant activity against lymphoid leu- kemia. This comprehensive report discussed the relationships of Camptotheca within the Nyssaceae, indicating that Camptotheca was closely related to Nyssa and only remotely related to Davidia. Research done by Titman (1949) using wood anatomy, Eyde (1963) using fruit structure and the fossil record, and Sohma (1963) using pollen support the taxonomic conclusions of Perdue et al. (1970). Our recent serological data also indicate that Camptotheca is more similar to Nyssa than to Davidia, and that Nyssa is more similar to Davidia than is Camptotheca (Tables 1-2). Our serological data are also supported in part by the findings of Hohn & Meinschein (1976) based on the fatty acid composition of seeds. They indi- cated that primitive Davidia and advanced Camptotheca were placed on each side of Nyssa, which is intermediate. Thus all the data presented lend credence 1977] FAIRBROTHERS—PLANT SEROTAXONOMY 157 to the placement of Camptotheca and Nyssa in the subfamily Nyssoideae and Davidia in the Davidioideae of the Nyssaceae. Serological experiments with four species of Nyssa have included compari- sons of Nyssa aquatica, N. biflora, N. ogeche, and N. sylvatica. The various experiments indicated that N. biflora and N. sylvatica were serologically very similar. Nyssa ogeche and N. aquatica were serologically distinct from each other and from N. biflora and N. sylvatica. The data also showed that sero- logically N. aquatica was more similar to N. biflora and N. sylvatica than to N. ogeche. However, N. ogeche was more similar to N. aquatica than to N. biflora and N. sylvatica. Nyssa ogeche was the most distinct species of the four com- pared (Fairbrothers & Johnson, 1964; Fairbrothers, 1966a, 1966b, 1968: Tables 1-2), The serological data support the conclusions of both Eyde (1963) and Sohma (1963), who reported close similarity between N. biflora and N. syl- vatica and treated them as two varieties of one species. The serological data does not support the findings of Hohn & Meinschein (1976) based on seed oil fatty acids. They indicated N. biflora and N. sylvatica to be chemically dis- tinguishable species. The various researchers agree that N. ogeche is the most distinct from the other three species of Nyssa. The serological data has not clearly indicated whether Camptotheca, Davidia, or Nyssa has the greatest similarity with Cornus. The three genera are serologically relatively similar to Cornus; however, the data indicate that Camptotheca might have slightly more similarity with Cornus than do Nyssa or Davidia (Tables 1-2 The genus Corokia (6 species) is restricted to the South Pacific region, rang- ing from northern New South Wales, Lord Howe Island, New Zealand, Chatham Islands, and Rapa Island, a distance of 4,000 miles. The serological data reveal very little similarity between Corokia cotoneaster and any species of the Cornaceae and Nyssaceae tested (Fairbrothers et al., 1975; Tables 1-2). Most researchers have indicated that this genus has little affinity with mem- bers of the Cornaceae in which it is often placed. Some botanists have suggested an affinity with the Saxifragaceae within the subfamily Escallonioideae (Escal- loniaceae) (Philipson, 1967; Smith, 1958). Eyde (1966, 1967) concluded that it was unrelated to Cornus but was possibly linked with Argophyllum. Kubitski 963) retained the genus in the Cornaceae. Hegnauer (1965) indicated that the Cornaceae may be related to either the Saxifragaceae or Loganiaceae. Takhta- jan (1969) excluded the genus Corokia from the Cornales and placed it in the Escalloniaceae (Saxifragales). Cronquist (1968) considered Corokia as a pos- sible nonmissing link between the Cornaceae and Escalloniaceae, Grossularia- ceae, or Saxifragaceae sensu lato. Both Cronquists and Takhtajan's classifica- tions reflect the serological data which indicate the distinctiveness of Corokia from members of the Cornales. However, Bate-Smith et al. (1975) investigated the distribution of several chemical compounds in the Cornales and concluded that Corokia possesses a chemical pattern consistent with that of the Cornaceae. The experimental investigation of taxa within the Cornales has indicated that the use of diverse disciplines has provided valuable data for helping to 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 understand the evolutionary development and relationships of the families and genera in the order. This order has proven to have been an excellent one for diverse chemosystematic research. Serological comparisons have provided signifi- cant data for evaluation in the continuing investigation of diverse cornaceous sensu lato, taxa. Taxonomy eventually must strive to bring together, summarize, and utilize what is known about the organisms to be compared. Systematic serologists have essentially learned to use the properties of one of the classes of proteins, gamma globulins, in comparative studies. Systematic serology provides comparisons which are relatively objective measurements; however, like all detected relation- ships, they are relative and not absolute. 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Ribulose-1,5- dub Me: carboxylase of Chlorella ellipsoidea. Arch. Biochem. eo 129: 597-602. TAKHTAJAN, A. 1969. Flowering Plants: Origin and Dispersal. Oliver & Boyd, Edinburgh. THORNE, R. F. un Synopsis of a putatively phylogenetic classification of the flowering plants. Aliso 6: 57—66. 1976. A phylogenetic classification of the angiospermae. Pp. 35-106, in M. C. UT W. C. Steere & B. Wallace (editors), Evolutionary Biology. Vol. 9. Plenum Press, New York Tirman, P. W. 1949. PH in the wood anatomy of the family Nyssaceae. J. Elisha Mitchell Sci. Soc. 65: 261. Urano, K. 1955. oa ae investigations on diagnosis of strain- relationship in co plant by precipitive tests I. Precipitive reaction between inbred lines of corn. Proc. adn Sci. Soc. Japan 24: 4-7. Vaucnan, J. G. & A. Warre. 1965. Taxonomic investigation of several Brassica species using serology and the separation of proteins by electrophoresis on acrylamide gels. Nature 208: 704-705. A. J. MacrEop & B. M. G. JONEs anes 1976. The Biology and Chemistry of the Cruciferae. Academic Press, London Wacerin, W. 1910. Nyssaceae. In A. Engler (editor), Das Pflazenreich IV. 220a, (Heft 41): 1-19. Wilhelm 5 (Ro Wncur, C. A. (editor). 1974. Biochemical and Immunological Taxonomy of Animals. Academic Press, London. ELECTROPHORETIC EVIDENCE AND PLANT SYSTEMATICS L. D. GOTTLIEB! ABSTRACT The study of phenotypes and their variation often provides evidence for phylogenetic in- ferences in plant systematics. Therefore, it is critical that the phenotypes analyzed reflect as directly as possible the underlying genotypes. The equation between phenotype and genotype is simpler and better understood for evidence obtained by electrophoresis of plant enzymes than for most morphological characters. This article riis the advantages and limitations of imo. yapana evidence to test hypotheses in plant systematics and evolution. It also sum- marizes the results of a large number of studies which have utilized this evidence. Three gen- eral 1 pe these studies are: (1). Conspecific plant populations are "ari similar genetically as pou by their very high mean genetic identities, 0.95 + 0.02. This result suggests that one a few populations often constitute an adequate sample of a species. 775 e sini species have strikingly reduced mean genetic identities, 0.67 + vever, certain pairs of annual plant species have genetic identities similar to those of pan i 5 In these cases, the species have been shown to be related as progenitor and derivative with the derivative being of recent origin. ( The amount of genetic variability within plant populations appears closely correlated with their breeding system, with outcrossing populations substantially more variable than inbreeding ones. The article 7 describes a number of actual and potential applications of electrophoresis in plant systematic Evidence obtained by electrophoresis of enzymes has not been widely utilized by plant systematists although it has dominated the research of many of their zoological poa po and population geneticists ( Manwell & Baker, 1970; Lewontin, 1974; 1975; Ayala, 1976). This has meant that the strengths and weaknesses of s eni for solving systematic and evolutionary questions in plant biology have not been sufficiently discussed. The present article is de- signed to facilitate an efficient evaluation, and emphasizes the unique charac- teristics of electrophoretic evidence, the requirements for its analysis, and actual and potential applications in plant systematics and evolution. ELECTROPHORETIC EVIDENCE; ADVANTAGES AND LIMITATIONS The systematist analyzes phenotypes and their variation and often uses this evidence for phylogenetic inference. Such inferences require that observed phenotypes have a specifiable relationship to unobserved genotypes. The equa- tion between phenotype and genotype is simpler and better understood for electrophoretic evidence than it is for evidence obtained from morphological characters or chromatographic comparisons of secondary metabolites. This fol- lows from the colinearity of amino acid sequence and nucleotide sequence as well as the specificity of enzyme catalysis. It also reflects the fact that electro- phoretic evidence is used to answer a very different kind of question than has usually been posed by systematists. kaysa as analysis answers a question such as: Are flower petals with Department of Genetics, University of California, Davis, California 95616. ANN. Missouni Bor. Garp. 64: 161-180. 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 conspicuously lobed limbs present in taxon A and taxon B? Chromatographic analysis asks, for example: Is apigenin present in both taxon A and taxon B? In contrast, electrophoresis answers a question of a different kind: Does gluta- mate dehydrogenase have the same electrophoretic mobility(ies) in taxon A and taxon B? Such a question probes the physical properties of particular en- zymes or other proteins on the hypothesis that these properties reveal, to a large degree, the record of accumulated mutations that have taken place in the gene specifying the enzyme. And that when a number of enzymes are considered simultaneously, this record can be evaluated with more precision and objectivity than highly complex morphological features. The primary observed evidence in studies of electrophoretic variation in natural populations is bands of color in a slab of starch or acrylamide gel. These mark the positions reached by different molecular forms of an enzyme (or other protein) which have migrated through the gel under the influence of an electric field. The enzyme variants are separated because they have different electrostatic charges (a function of the relative number of amino acids with positive and nega- tive charges on their surface). Their migration through the gel may also be dif- ferentially affected by their size or configuration. Following their physical sepa- ration, the enzymes are identified by a staining reaction based on their catalytic activities. The combination of electrophoresis and staining specificity makes it possible to distinguish particular enzymes among hundreds that may be present in a crude tissue extract. The different molecular forms of an enzyme that catalyze the same reaction are called isozymes if their polypeptide constituents are coded by more than one gene locus (e.g. lactate dehydrogenase, human ADH). They are called allo- zymes if their polypeptides are specified by different alleles at a single gene locus; the majority of enzymes routinely studied in natural populations have different allozymic forms. Allozymes are the biochemical consequence of the substitution, deletion, or addition of amino acids in the polypeptides which comprise the enzyme, and they can be distinguished if these changes affect their electrophoretic migration. Since the amino acid sequence of a polypeptide is colinear to the nucleotide sequence of its coding structural gene locus, allozymes result from gene mutation. Thus, an analysis of protein structure using electrophoresis is, to a first approxima- tion, an analysis of a gene. It is precisely this simple relationship between the bands of color on the gel and the nucleotide sequence of genes that makes pro- tein electrophoresis a powerful analytic tool for systematics. major advantage of electrophoretic evidence is that colinearity assures that systematic comparisons can be made between products of genes which are homologous (have a common origin), thus, avoiding problems of convergence and functional correlation often prevalent with morphological characters. An- other important advantage is that electrophoretic evidence is precise and di- rectly quantifiable in terms of the number and kinds of enzymes studied, per- mitting the amount of genetic information utilized to be stated exactly. This is seldom possible with morphological or other characters. A third significant advantage is that comparisons are made with enzymes that are generally always 1977] GOTTLIEB—ELECTROPHORESIS AND PLANT SYSTEMATICS 163 present (with the exception of those selectively turned on or off during develop- ment) and little influenced by environmental factors. This avoids the frequent situation that occurs with both morphological and chromatographic data in which the absence of a character in one taxon is interpreted as an indication of a less close phylogenetic relationship between it and other taxa that display the charac- ter even though independent evidence is lacking regarding the cause of the charac- ters absence. Another advantage is that problems of a priori character weighting do not occur with electrophoretic evidence because all enzymes examined are accorded equal value in similarity matrices or other methods of evaluating di- vergence. The theoretical advantages of electrophoretic evidence, to be sure, are offset by certain shortcomings but, fortunately, these are reasonably well defined. The first problem is that a small number of enzymes is sampled and these may not represent enzymes in general since they are most often involved in some aspect of glycolysis, intermediary metabolism, or in the catalysis of certain general types of bonds (esterases, phosphatases, peptidases). Lack of representativeness is probably less serious for systematics, which has not confined itself to charac- ters thought to be representative, than it is for genetic studies which attempt to estimate the total amount and kind of genetic variation in different kinds of or- ganisms. In any event, the enzymes that are examined comprise a sufficiently large category to give meaningful information about many kinds of evolutionary changes. A problem which is more critical for systematics is that, even for those en- zymes examined, the redundancy of the genetic code means that only about 30% of the substitutions of nucleotides are expected to result in the substitution of amino acids that cause changes in electrophoretic mobility (Shaw, 1970). In addition, allozymes that have identical mobilities do not necessarily have identi- ‘al amino acid sequences. In fact, recent studies utilizing amino acid sequenc- ing (Boyer et al., 1972), heat denaturation (Bernstein et al., 1973; Singh et al., 1974), and variation in gel pore size (Johnson, 1976) suggest that a single mo- bility class on a gel may sometimes contain more than one enzyme. This requires that more weight be given to evidence of electrophoretic difference than to evi- dence of similarity. Additional biochemical tests, however, are available to de- termine whether allozymes with the same mobility have different amino acid sequences. All in all, electrophoretic evidence should be regarded as providing, an underestimate of the actual amount of genetic difference between taxa. Electrophoretic evidence does not include any information on the number of amino acid differences, or mutational steps, that cause differences in enzyme mobilities. A difference in mobility can reflect a single nucleotide substitution or numerous changes in nucleotide sequence. Thus, electrophoresis can demon- strate that two taxa have different allozymes of phosphoglucoisomerase, but it does not provide information about the amount of difference. This is likely to be greater with increasing phylogenetic distance. Thus, the equation between phenotype and genotype remains acceptable even when the limitations of electrophoretic evidence are considered because they can be specified and the direction of bias is generally known. 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ELECTROPHORETIC EVIDENCE: A FRAMEWORK FOR ANALYSIS When tissue extracts are subjected to electrophoresis in a starch gel, the pat- tern of enzyme bands (number, spacing, and intensity) is an expression of the particular enzyme system assayed and its mode of inheritance. For some en- zymes (usually esterases, phosphatases, and peroxidases), single individuals dis- play complex patterns with as many as 10 to 15 bands because they possess numerous gene loci that code different molecular forms. In contrast, other en- zymes are specified by a single structural gene and individuals might display only a single band following electrophoresis. The number of polypeptide sub- units of each enzyme and the allelic state of the coding gene (homozygous or heterozygous) also determine the number of enzyme bands displayed. Thus, for an enzyme composed of a single polypeptide, an individual heterozygous at the coding gene displays two allozyme bands, but if the enzyme is dimeric (composed of two polypeptides), three allozymes are displayed, and if it is tetrameric, five are displayed (Fig. 1). In other cases, the polypeptide constit- uents of an enzyme are coded by different nonallelic genes, for example, alco- hol dehydrogenase in maize (Freeling & Schwartz, 1973) and in sunflower (Tor- res, 1976), producing still additional variants. The number of enzyme bands can be reduced if enzymes specified by dif- ferent genes overlap on the gel because they have similar mobilities, or if an individual is homozygous for a “null” allele (an absence of activity which genetic analysis demonstrates to be allelic to genes that specify active forms of the en- zyme). Artifacts that might result from procedures of extraction or electrophore- sis can also change band number. In addition to these biochemical factors, the pattern of enzyme bands displayed by different individuals in a population is a function of the amount of genetic variation for the enzyme system. The presence of so many factors which influence the appearance of the electro- phoretic phenotype means that the systematist must reject the temptation to com- pare electrophoretic data from different taxa by direct inspection, i.e., simply counting the number of bands with similar and dissimilar mobilities. Not only would this approach be biochemically faulty, but it nullifies several of the im- portant advantages of electrophoresis, particularly the inference of enzyme homology, and the precise and quantifiable form of the evidence. The frequent complexity of the electrophoretic phenotype means that, at least for the complex systems that have a number of electrophoretically separable enzyme variants, a genetic analysis is necessary. Such analysis demonstrates which variant forms of an enzyme system are specified by allelic genes and which by nonallelic genes; i.e., it distinguishes allozymes and isozymes. In addition, in many cases, it rules out the possibility of biochemical artifacts. Genetic analysis also leads directly to a quantitative speci- fication of the electrophoretic data that usually is ordered as follows: the num- ber of structural genes specifying the enzymes examined; the proportion of genes that show variation or, as the geneticist says, are polymorphic in that they have more than a single allele; the number of alleles per gene in the population; and the mean proportion of genes which is heterozygous per individual. The protocol of formal genetic analysis can often be simplified with electro- 1977] GOTTLIEB—ELECTROPHORESIS AND PLANT SYSTEMATICS 165 These diagrams present three examples of electrophoretic patterns in parents and their hybrids to illustrate that differences between Tama ay. in the number of enzyme bands often do not equal the number of their genetic nao Jase 1 presents a cross be- tween two individuals, homozygous I different alleles at a gene 1 : an enzyme. The num- ber of bands per individual in their F ybrids depends on the subunit structure of the enzyme. Note that for a dimeric enzyme, a Ran cn individual differs from each of its par- ents by two bands, but one allele; for a tetrameric enzyme, it differs from them by four bands, but one allele. Case 2 illustrates the same point but uses a cross between individuals homozy- gous for the same allele at one gene and homozygous for different alleles at a second gene. The F; phenotype depends on the subunit structure of the enzyme. In this example, poly- peptides specified by the two genes do not have affinity for one another. Case 3 presents a m e = two individuals whose enzyme phenotype is also determined by two gene loci. | this case, the enzymes are considered to be dimeric, and polypeptides specified by both prd “alleles a different genes associate to form "hybrid" or heteromeric enzymes. The cross is a test cross between a double heterozygote and a aed q V If the genes assort independently, ied ie progeny classes are produced in Pune if they are linked, the parental phenotypes are more frequent than the den: 2 n Note that the double heterozygote ditfers "n the double homozygote by six bands 125 m one allele at each gen 1e polypeptide structure of each enzyme band is given on the left and the geno- type of esoh individual is given below. CASE 1 Number polypeptides in enzymes 1 2 4 x — m — Parents F, phenotypes CASE2 E B x — — — Parents F, phenotypes CASE 3 RF lala = — lalb —— EI — lblb — — —— — — — 1422 — — — la2b + 1b2a— X — — — — — — lb2b —— —— —— 2a2a ——— — COME aa — — 2a2b —— — — 2b2b —— — — la 2a Ib 2a la 2a 1b 2a Ib 2a la 2a lb 2b lb 2a lb 2b Ib 2a lb 2b Ib 2a Parents Progeny phenotypes phoretic data because enzyme bands, with few exceptions, show codominant in- heritance and segregate as single Mendelian factors (see reviews by Scandalios, 1969, 1974; Jacobs, 1975). This makes it possible to utilize progeny tests to re- place formal crosses between individuals with different phenotypes, examine Fy progeny phenotypes, and analyze phenotypic segregation patterns in the F, 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 generation. In the progeny tests, individuals are grown from open-pollinated seeds collected on single plants in nature. Since these individuals all have one parent in common, they necessarily have in common its alleles. For many enzyme systems, study of the segregation pattern of the different variants in such prog- enies can indicate which of them are specified by allelic genes and which by different gene loci (Brown et al., 1975). However, for very complex systems in which several polymorphic genes and overlaps in the migration of different en- zymes are involved, formal analysis is still required. Figure 1 presents examples of such analysis. Before the introduction of the electrophoretic technique, the study of genetic variation in natural populations was unsatisfactory because it depended on the identification and enumeration of rare recessive mutants that, when homozygous, yielded visible morphological changes. The genetic basis of many of these charac- ters is simple and clearly demonstrable, but they constitute only a very small proportion of the genetic variation in populations. The vast majority of pheno- typic characters are apparently controlled by many genes each of which may have different individual effects. These so-called quantitative characters are also often strongly influenced by environmental variation. The result is that the contribution of individual genes of this type cannot be ascertained, and the extent to which they vary one from another in different individuals is undetectable. Thus, one studied either those rare characters controlled by one or two genes with major effects or the much more common characters controlled by many genes that are neither individually identifiable nor distinguishable from environmental in- fluences. In consequence, the traditional methods of studying genetic variation were stymied by the impossibility of equating phenotypes with genotypes. The electrophoretic procedure avoids most of these problems. In addition, it identifies genes which do not vary on the basis that their enzyme products do not vary in their electrophoretic mobility (within the limits mentioned in the previous section), making it possible to determine the proportion of genes that show variation. This advantage has been considered the “cornerstone” of the method (Hubby & Lewontin, 1966) because previously it was not possible to equate lack of variation with monomorphism at particular genes. The determination of how many individuals and populations to sample before a confident statement can be made regarding the amount of genetic divergence between taxa is another important consideration in the use of electrophoretic evi- dence. A definition of the meaning of divergence greatly simplifies this prob- lem. Thus, maximal divergence between two taxa at a gene locus means that they have no alleles in common. Minimal divergence at a gene locus means that the two taxa have similar complements of alleles in similar frequencies. Although the two extremes are connected by a wide variety of intermediate situations, the amount of divergence at a set of genes can often be fitted into a general pic- ture. Thus, if a large number of genes is studied, about 30-50% of them are likely to be monomorphic, another 30-40% will be moderately polymorphic (two or three alleles), and the remaining genes will be highly polymorphic. The distribution of genes in monomorphic and polymorphic categories per- mits the systematist to decide, at the outset of a study, the amount of sampling 1977] GOTTLIEB—-ELECTROPHORESIS AND PLANT SYSTEMATICS 167 necessary to test a given hypothesis. For example, at a polymorphic gene locus, an allele can be considered common (moderate to high frequency) and wide- spread, rare (less than 0.05) and widespread, common and local (one or two populations), or rare and local. For certain systematic purposes, one might de- cide that the first category is most relevant (it is the easiest to sample since al- leles here have the highest probability of being included in a sample regardless of strategy). In logistic terms, this means that relatively little effort need be ex- pended to find one more allele which is likely to have low frequency, be local in distribution, or both. The number of individuals to sample per population is best viewed from the standpoint of how many plants to examine in order to have a 95% certainty of observing all the alleles at a locus which have frequencies greater than 0.05. This problem has been considered by Marshall & Brown (1975) who show that, even in the unlikely case of 20 alleles with frequencies of 0.05 each, a random sample of 120 gametes (60 individuals) will include, with 95% certainty, one copy of each allele. In sum, decisions related to sampling can be neatly bracketed be- cause electrophoretic evidence consists of discrete and precise units of informa- tion. A number of coefficients have been developed to summarize allele frequency data into a single figure that might be used to assess the degree of genetic di- vergence of taxa (Cavalli-Sforza & Edwards, 1967; Hedrick, 1971; Nei, 1972: Rogers, 1972); however, all of them appear to provide similar estimates (Avise, 1974). The data can also be used to construct dendrograms that cluster taxa ac- cording to their similarities (references in Avise, 1974). Another approach, de- scribed in the following section, that might be particularly useful for analysis of conspecific populations, makes use of the presence or absence of alleles rather than their frequencies ( Gottlieb, 1975). ELECTROPHORETIC EVIDENCE: APPLICATIONS Electrophoretic analysis of enzyme variation provides efficient, quantitative estimates of the amount of genic variation within natural populations and the ex- tent of genic divergence among populations. A very large number of electro- phoretic studies have been made on animal species. The results appear remark- ably consistent: (1) Single populations of both vertebrates and invertebrates contain substantial genetic variability, and perhaps as much as 90% of the total genetic information of their species (review in Powell, 1975; Selander, 1976); (2) Conspecific populations have a very high degree of genic identity (Nei, 1972), often with a mean above 0.90, on a scale of 0 to 1. Their high identity reflects the fact that the same allele is usually fixed at monomorphic genes (as much as 80 to 90% of the genes in vertebrates, for example), and, at the poly- morphic genes, only the frequency of alleles differs (Avise, 1974); (3) Closely related species are considerably more differentiated than conspecific popula- tions, with a mean genetic identity around 0.50 to 0.60, which suggests that dif- ferent species of animals are almost completely distinct in allelic composition at about one-quarter to one-half of their genes (Ayala, 1: . Many of these differences may have evolved since their origins as species Ede much higher 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 values of genetic identity are observed for pairs of species which apparently originated in the Pleistocene (Nevo et al., 1974; Avise et al., 1975). In contrast to the wealth of data for animal species, the number of electro- phoretic studies of plant species is extremely small (fewer than a dozen groups of congeneric species have been examined for electrophoretic variation in a large number of enzymes) and, consequently, generalizations must still be considered tentative. The paucity of studies with plants is unfortunate because, in many ways, plants are better material than animals for electrophoretic investigations. Thus, they are often easy to grow in high numbers; they need not be killed to obtain a tissue sample so that individuals can be used for additional analysis; progeny-testing to establish the genetic control of enzyme variants is straight- forward; natural populations are often spatially and ecologically delimited, per- mitting coordinated studies of ecological adaptations and amplitudes; phylogenies are often known unambiguously, facilitating analysis of the consequences of speciation; breeding systems are highly variable so that genetic consequences of different amounts of inbreeding can be studied directly and correlated with demographic inputs, etc. The available electrophoretic studies with plants that are relevant to ques- tions in systematics and evolution can be grouped into four major categories: (1) genetic divergence among conspecific populations; (2) genetic divergence among congeneric species, a subject which has also provided evidence of the genetic and biochemical consequences of speciation; (3) enzyme expression in diploid progenitors and polyploid derivatives; and (4) a heterogeneous group of special-purpose studies (not reviewed here because of space limitations) deal- ing with themes such as analysis of gene flow across species barriers (interspecific hybridization) ( Chu & Oka, 1970; Levin, 1975), consequences of unusual chromo- somal pairing mechanisms in Oenothera (Levy & Levin, 1975), demographic analysis (Schaal, 1975), the effect of breeding systems on the amount and ex- pression of genetic variability ( Allard, 1975; Allard & Kahler, 1971), and genetic diversity and edaphic specialization ( Babbel & Selander, 1974). ELECTROPHORETIC EVIDENCE: CONSPECIFIC POPULATIONS Electrophoretic variation in enzymes has been examined in natural popula- tions (at least two) of about 28 plant species (Table 1). However, in many re- spects, the data is very uneven. For example, the number of enzyme systems examined and the number of genes that code them in the different species vary over a five-fold range. In addition, the choice of enzymes is diverse so that in some studies the proportion of genes specifying enzymes that are frequently highly polymorphic (esterases, phosphatases, peroxidases) is high, whereas in other studies many additional enzymes are included that are involved in basic metabolism (such as phosphoglucoisomerase, phosphoglucomutase, glutamate de- hydrogenase, malate dehydrogenase, malic enzyme, glutamate oxaloacetate trans- aminase). Further, the number of populations sampled per species varies widely. 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DIYADIO 6261 ‘PIY X 38900 FSO €0'0 00'S 001 S/E L «AIL, Dipqapq `y €L61 ‘PNV X 3390 001 00°0 = 00°0 987 6 valn qq nuoay aoualajay A1quop AMSOSÁZOI9]9H Əəuə5 ul o1qdaouiAqoq səuə9 ‘ON ‘dog ‘ON səroədç houses uvoN /S9[9II V sauay % /sui91sAg ON uv oN 9uiÁzu^q ‘ON 'suoge[ndod om} jsee[ ye ur pəzÁjeue sem səuiKzuə jo 19quinu egie| € YY ur sorods [uo sopn[our sı səəds au "eue ? se powon safoads done 10j Ə payussaid ejep oq -suone[ndod jue[d orjroəds -uoo JO (¿61 “ION JO nens OY} YWA poje[no[e?) sonmnuopr onauos pue souikKzuo SurKgoods səuəñ 10j uonervA oneaioqgdonoopq 'I Adv 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 dominant selfers, and recently evolved and ancient taxa. However, in spite of these disparities, two results appear well established. First, the amount of genetic variability within plant populations is closely correlated with their breeding systems (Table 1). Thus, the mean number of alleles per polymorphic gene averages 1.88 + 0.12 for highly self-pollinating species and 2.86 + 0.24 for outcrossing ones. The mean proportion of genes that is heterozygous per individual follows suit: 0.032 + 0.013 for selfers and 0.133 + 0.026 for outcrossers. These results indicate that breeding system influences not only the degree of genetic homozygosity but also the total amount of vari- ability that can be maintained in plant populations. Second, conspecific plant populations are extremely similar genetically as demonstrated by their very high mean genetic identity 1 = 0.95 + 0.02, averaged over all species (Table 1). This is an important result for systematics because it suggests that electrophoretic evidence from one or a few populations very often constitutes an adequate sample of an entire species. e very high degree of genetic similarity among conspecific populations leads to the suggestion that should populations be discovered which have novel alleles or distinct allele frequencies at more than a few gene loci, such popula- tions are very likely to constitute distinct taxa and should be further examined with this in mind. Such evidence has already been used to identify a subspecies of Drosophila willistoni (Ayala, 1973) and a new species of sea cucumber (Man- well & Baker, 1963). It is not unlikely that electrophoretic evidence will also be similarly used to identify hitherto unrecognized plant species. An approach to the comparison of conspecific populations that considers the representativeness of single populations rather than their identity to one another has also been proposed (Gottlieb, 1975). Designated the Complement Index, it compares the number of nonunique and nonubiquitous alleles (present in more than one but not all populations) in each population with the total number of such alleles identified in all the populations examined. Since the presence of alleles increases the biochemical repertoire of a population, the number of alleles constitutes a very good and easily obtained estimator of the relative po- tential of different populations for adaptive evolutionary change. The Comple- ment Index would be particularly useful for taxa in which populations contain large numbers of different low frequency alleles, a situation that may be com- mon in outcrossing plants. Thus, 11 populations of Stephanomeria exigua subsp. carotifera all possessed the same gene at 6 monomorphic loci and 13 high fre- quency alleles at 8 polymorphic ones, but different numbers of 25 other low frequency alleles (Gottlieb, 1975). Calculation of the Complement Index showed that the populations actually represented subsp. carotifera to very different de- grees even though they had a mean genetic identity, J = 0.98 (Gottlieb, 1975). The average population had only about half of all the genes identified in the subspecies as a whole; nevertheless, one of them possessed every one of the non- unique alleles. This population contained more of the genetic resources of subsp. carotifera than any other and is the most likely to persist through environmental fluctuations. In addition to identifying such populations (which, with cultivated plants have obvious importance for germ plasm conservation), the Complement 1977] GOTTLIEB-—ELECTROPHORESIS AND PLANT SYSTEMATICS 171 Index could be used to compare the representativeness of populations of dif- ferent taxa. ELECTROPHORETIC EVIDENCE: CONGENERIC SPECIES The large amount of electrophoretic variation in natural populations of plants initially led systematists to presume that extensive surveys within and between populations were required in order to use electrophoretic evidence for meaning- ful systematic comparisons between species (Turner, 1969). This point of view apparently took hold because genetic studies had only rarely been carried out and therefore it was difficult, if not impossible, to make sense of the complex banding patterns that were observed (many of the early studies unwittingly utilized esterases and peroxidases which are the most difficult systems to inter- pret). The absence of genetic data and the small number of populations that had been sampled combined to give the impression that the polymorphisms in- herent in electrophoretic evidence lessened its value for systematics. However, now it is realized that once polymorphisms are defined in genetic terms so that “bands” can be equated with alleles and different gene loci, then their presence actually increases the power of electrophoretic evidence for sys- tematic studies since they reveal to a large degree the accumulated record of nu- merous mutations that have become established in the coding genes. In addition, extensive sampling of conspecific populations is often not necessary for species comparisons because many of the alleles, particularly those with frequencies above 0.20, as well as the genes at monomorphic loci, are now known to be present in most, if not all, populations of a species. However, although the number of populations sampled can be reduced, it remains important to increase the num- ber of enzymes sampled. This would tend to lessen the effect of biases that might result from selecting only enzymes likely to be polymorphic or those limited to any particular biochemical category. About a dozen studies have been made that provide evidence of the extent of genetic divergence between congeneric species ( Table 2). These studies show that most pairs of species have strikingly reduced genetic identities; J = + 0.07, averaged over all pairs of species examined. A number of species pairs, however, have very high genetic identities, within the range of those characteris- tic of conspecific populations. For three of these cases, Stephanomeria exigua subsp. coronaria and “Malheurensis” (Gottlieb, 1973b, 1976), Clarkia biloba and C. lingulata (Gottlieb, 1974a), and Gaura longiflora and G. demareei ( Gott- lieb & Pilz, 1976), the species are known to be related as progenitor and deriva- tive, respectively, with the derivative being of relatively recent origin. e three cases represent the three possible pathways of diploid speciation in annual plants, defined in terms of the breeding systems: self-incompatible to self-compatible, self-compatible to self-compatible, and self-incompatible to self-incompatible, respectively. A fourth example of very high genetic similarity between progenitor and derivative diploid species has recently been identified in Lycopersicon ( Rick et al., 1976). The lack of genetic divergence between the members of each species pair indicates that, shortly after their origin, annual plants, regardless of their breeding system, are still limited genetic versions of 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE 2. Neis (1972) mean genetic identity, I, between pairs of populations of con- generic plant species. Species I Reference Clarkia biloba X C. lingulata 0.88 Gottlieb, 9 80 Clarkia rubicunda X C. franciscana 0.28 Gottlieb, Gaura longiflora X G. demareei 0.99 Gottlieb : po 1976 Hymenopappus scabrosaeus X rtemisiaefolius 0.90 Babbel & Selander, 1974 Lupinus texensis X L. subcarnosus 0.35 Babbel & Selander, 1974 Oenothera strigosa X O. biennis 0.97 evy & Levin, Oenothera strigosa x O. parviflora 0.54 Levy & Levin, 1975 Oenothera biennis x O. parviflora 0.55 Levy & Levin, 1975 hlox drummondii x P. cuspidata 0.67 Levin, 1975 Stephanomeria exigua subsp. coronaria Malheurensis" 0.94 d 1973b, 1976 Tragopogon dubius X T. porrifolius 0.50 e & Gottlieb, 1976 Tragopogon dubius X T. pratensis 0.62 1 12 5 & Gottlieb, 1976 Tragopogon porrifolius >x T. pratensis 0.53 Roose & Gottlieb, 1976 their progenitors and possess very few or no unique alleles insofar as their genomes have been assayed. The species are extracted from the parental repertoire of phenotypic variation and genetic polymorphisms. Thus, the speciation process does not seem to involve early reconstitution of the genome of the derivative species, even though it may possess certain unique morphological traits or other features ( Gottlieb, 1976). The other two examples of high genetic identity between species are not in- consistent with this thinking. Thus, the similarity of Oenothera strigosa and O. biennis presumably reflects the fact that they have one genome in common (Levy & Levin, 1975). And the high identity of the two species of Hymenopappus is concordant with their very close phylogenetic relationship as judged by their high overall morphological similarity; they were maintained as species because they apparently do not hybridize despite extensive parapatric contact (Turner, 1956). However, it is not clear if the results with annual plants will also characterize perennial plant species. This is because perennials, especially long-lived ones, appear to evolve gradually, rather than rapidly and abruptly, and they are more likely to be reproductively isolated by ecological and pollination factors rather than hybrid sterility resulting from chromosomal restructuring. Electrophoretic evidence has also been used to show that species which ap- pear similar may actually not be so. Thus, Clarkia franciscana, a highly self- pollinating species thought to have evolved by rapid reorganization of chromo- somes from the morphologically similar C. rubicunda (Lewis & Raven, 1958), is totally divergent from that species in a high proportion of its genes (those coding six of the eight enzyme systems assayed) (Gottlieb, 1973a). In addition, C. franciscana has a duplicated gene for alcohol dehydrogenase which further distinguishes it from C. rubicunda (Gottlieb, 1974b). Such marked genetic dif- ferentiation requires that the phylogenetic separation of the two species oc- curred much longer ago than had been presumed, and makes its proposed mode of origin quite uncertain. Therefore, a reasonable criterion to apply in cases like 1977] GOTTLIEB ELECTROPHORESIS AND PLANT SYSTEMATICS 173 this is that a species not be accepted as having originated recently from another extant species if it is not electrophoretically highly similar to its putative parent (Gottlieb, 1973a). This criterion has now been satisfied in the four examples de- scribed above. When additional electrophoretic studies are reported, it may very well turn out that such evidence reflects species divergence more sensitively than other types of biochemical analysis. Thus, two-dimensional chromatography of cer- tain flavonoids in Tragopogon dubius, T. porrifolius, and T. pratensis, failed to distinguish a single component that was species-specific (Brehm & Ownbey, 1965), even though the morphological differences between these species are "broad, sharp and absolute" (Ownbey, 1950). But, electrophoretic analysis of many enzymes in North American populations of the three Tragopogons re- vealed very clearly that they were fixed for different alleles at about 40% of the 2] genes examined (Roose & Gottlieb, 1976). Other groups of plants have not yet been studied so extensively both for electrophoretic variation in enzymes and chromatographic variation in flavonoids and, therefore, it is not possible to know if such a result will prove general. However, this is not implausible since changes in the amino acid sequences of a large number of polypeptides which affect electrophoretic mobilities of enzymes are more likely to reflect early stages of genetic divergence than are changes in secondary metabolites such as flavonoids which are products of enzyme-catalyzed biosyntheses. Electrophoretic evidence is also likely to be useful in other systematic in- vestigations which require knowledge of the extent of genetic similarity of closely related diploid species. An attractive use will be to examine cases in which one species appears to be a stabilized derivative of hybridization between two other extant species such as Lasthenia burkei (Ornduff, 1976), Potentilla glandulosa subsp. hansenii (Clausen et al., 1940), Achillea rosea-alba (Ehrendorfer, 1959), and Delphinium gypsophilum (Lewis & Epling, 1959). Another use will be to answer a novel systematic question having to do with the relative similarity of species in different genera. A sample question might be: Are species of Clarkia more similar to one another than species of Baptisia? Once again the question becomes plausible because of enzyme homology and because electrophoretic evidence is composed of discrete, quantifiable units of information. The com- parison of relative taxonomic distance in different genera might eventually lead to the development of procedures to standardize certain taxonomic decisions. A further application of electrophoretic evidence above the species level takes advantage of its ability to distinguish species with different numbers of genes specifying the same enzyme system. In cases where an enzyme is composed of several polypeptides, the formation of "hybrid" enzymes by the association of subunits coded by different gene loci provides strong evidence for their homology and the origin of one of the genes through duplication. Clearcut examples of such gene duplication have been documented for animal lactate dehydrogenase (reviewed in Markert et al., 1975), hemoglobins (Ingram, 1961), and phospho- glucoisomerase ( Avise & Kitto, 1973). Gene duplication is very probably a unique event in the evolutionary history of organisms and, therefore, it can provide evidence of the monophyletic origin of large groups of species and genera. 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 CLARKIA PGI EUCHARIDIUM GODETIA C C. WILLIAMSONIL PHAEOSTOMA PRIMIGENIA C. XANTIANA C. AMOENA PERIPETASMA C. FRANCISCANA C. BILOBA C. RUBICUNDA C. DUDLEYANA 2P G I GENES 3 P G Í GENES PGI-1— —PGI-1 A/A NA — — PGI 2 PGI-2 AB — 99 = — N BLIIEE Fi Eight diploid zo in five of the seven diploid sections of Clarkia which endi been examined to date can be ed into two groups on the basis of the number of genes they have specifying phosphoglucoisomerase subunits. The photographs show typical fee phoretic phenotypes for this enzyme system in the two groups. Details of the genetic analysis are described by € Gottlieb (1977). During studies of the genetic divergence of diploid species of Clarkia, I un- covered two cases of apparent gene duplication (Gottlieb, 1974b, 1977). The one involving duplicated phosphoglucoisomerase (PGI) is particularly relevant to systematic studies because it tests directly the taxonomic delimitation of sec- tions within the genus proposed by Lewis & Lewis (1955). Genetic analysis of PGI in Clarkia has shown that species either have two or three genes specifying these enzymes (Gottlieb, 1976, 1977). To date, eight species in five of the seven diploid sections of the genus have been examined: Clarkia rubicunda, C. amoena, and C. franciscana in the presumed primitive section Primigenia have two genes (Gottlieb, 1973a) as does C. williamsonii in the Godetia section (Price, 1975). Section Phaeostoma is represented by C. xantiana, Peripetasma by C. biloba and C. dudleyana, and Eucharidium by C. concinna, and all of these species have three PGI genes (Gottlieb, 1977) (Fig. 2). The duplication was originally recognized because individuals from species with three PGI genes display more enzymes upon electrophoresis than those from species with two PGI genes. The actual number of enzyme bands ob- served depends on the allelic state of the coding genes and the affinity of poly- peptides specified by the different alleles and the different gene loci. PGI is composed of two polypeptides so that an individual heterozygous for different al- leles at a single coding locus normally produces three enzymes by two-by-two association of the two polypeptides (aa, bb, ab). In the Clarkias examined, the 1977] GOTTLIEB—ELECTROPHORESIS AND PLANT SYSTEMATICS 175 gene coding the most anodal PCI appears to be invariant since individuals with either two or three PGI genes have always possessed a single fast PGI; poly- peptides specified by this gene, called PGI-1, do not form a “hybrid” enzyme with those specified by either of the other genes ( Fig. 2 us, the maximum number of PGI enzymes observed in the two-gene spe- cies was four (the single fast PGI coded by PGI-1 plus three enzymes in indi- viduals heterozygous at PGI-2). However, as many as ten enzymes have been observed in the three-gene species because polypeptides specified by the du- plicated gene, called PGI-3, from "hybrid" enzymes with those specified by the original PGI-2 gene. Thus, when both PGI-2 and PGI-3 are heterozygous, four different polypeptides are made which aggregate to form nine distinguishable enzymes (Fig. 1, case 3), and PGI-1 codes a tenth enzyme band. The duplication is thought to have originated by the generation of a dupli- cated chromosome segment in a progeny of a cross between individuals differing for chromosomal rearrangements, possibly a partially overlapping reciprocal translocation (reviewed in Burnham, 1962). Self-fertilization would make the segment homozygous in a few generations. This mode of duplication is likely in Clarkia because in this genus, species are self-compatible and differ by large numbers of reciprocal translocations. Such duplications will not be linked; recent genetic analysis (Gottlieb, 1977) in Clarkia xantiana has shown that PGI-2 and PGI-3 assort independently, which is consistent with the proposed duplication process. he species with three PGI genes can be considered a monophyletic group that traces back to an ancestor that branched away from the Primigenia-Godetia stock. That two genes is the ancestral number is directly supported by the ob- servation that Oenothera, the most closely related genus to Clarkia, also has two genes for PGI ( Levy et al., 1975) as does Gaura (Gottlieb & Pilz, 1976), another genus in the same tribe of the Onagraceae. The utilization of the number of genes coding specific enzymes to classify groups of species into monophyletic assemblages appears not to have a parallel in current systematic research. Gene duplication provides a strict homology, absent with most morphological charac- ters, because convergence in particular structural genes is highly improbable since it would require a very high number of mutational changes to alter the cod- ing properties of a different nonhomologous locus. ELECTROPHORETIC EVIDENCE: POLYPLOID SPECIES The ancestry of most allotetraploid species (tetraploids are used as an exam- ple of polyploids) can, in principle, be traced back to a chromosome doubling in a diploid individual which was produced by hybridization between differentially adapted populations. The initial allelic composition of the tetraploid plants is a direct function of the degree of genetic divergence of the diploid progenitor populations and is likely to be substantially greater if these represent species. This follows because species have a very much higher probability than con- specific populations of possessing different alleles at their monomorphic gene loci and nonoverlapping complements of alleles at polymorphic genes. After the events of its origin, the courses of evolution in the tetraploid and its diploid 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 parents are independent which suggests that the more ancient the tetraploid, the less likely will it retain the alleles it inherited in an unmutated state, and, likewise, alleles will continue to evolve in the diploids. Thus, the ability to identify the diploid parents of a tetraploid species with electrophoretic evidence depends on numerous factors having to do with the amount of divergence of the diploids at the time of tetraploid origin as well as subsequent evolutionary events. It follows that evidence brought to bear on the phylogeny of a tetraploid spe- cies not be limited to a few enzymes or to a limited class of proteins, and that particular attention be paid to specific protein homology and the mode of in- heritance of the proteins examined in order that an absence or difference in mobility can be interpreted in terms of genetic changes. This prescription has been ignored in numerous studies of diploid and tetraploid plant species which have employed only one or two enzyme systems or have sampled only seed pro- teins and, consequently, many of these studies have uncertain value and are not dealt with here. In general, electrophoretic analysis has demonstrated that polyploid species express, additively, enzymes present separately in their diploid parents. This result has been reported, for example, in wheat (Hart, 1969; Mitra & Bhatia, 1971; Barber, 1970), cotton ( Cherry et al., 1972), Nicotiana (Smith et al., 1970; Reddy & Garber, 1971; Sheen, 1972), Phaseolus (Garber, 1974), Stephanomeria (Gottlieb, 1973c), and Tragopogon (Roose & Gottlieb, 1976). If the duplicated genes of polyploids specify different polypeptide subunits of multimeric en- zymes (those that are composed of more than one polypeptide), additional “hy- brid" enzymes are produced which are not expressed in a diploid parent if it lacks both coding alleles. For many enzymes, the polyploid species is a "fixed heterozygote" because all of its individuals express a multiple enzyme phenotype that reflects their possession of different coding alleles inherited from the diploid species. This multiple enzyme phenotype does not exhibit genetic segregation because, at meiosis, chromosome homologues often pair preferentially so that genes inherited from both diploid parents go to the same pole and each gamete receives one copy of each of them. At fertilization, each gene is made homozy- gous, but their presence in duplicate means that a heterozygous ( multi-enzyme) phenotype can be produced in the tetraploid. The multiplicity of enzymes in a polyploid species may extend the range of environments in which normal de- velopment can take place, and this is a reasonable hypothesis to account for the frequent wider distribution of tetraploid species relative to the diploids in many genera (Barber, 1970; Manwell & Baker, 1970; Gottlieb, 1976). The general observation that the enzymes usually assessed by electrophore- sis are expressed additively in polyploid species (the only apparent exception is a study in wheat, Sing & Brewer, 1969) may reflect, to some extent, the rela- tively recent origin of the tetraploids which have been examined. Although an- cient polyploid complexes have not yet been studied by electrophoresis, many recently evolved polyploids have considerable systematic and evolutionary signifi- cance because they provide critical evidence regarding the initial genetic and bio- chemical consequences of this type of genome doubling. The most extensive comparison of enzyme variation in diploid and tetraploid 1977] GOTTLIEB—ELECTROPHORESIS AND PLANT SYSTEMATICS 177 species has been made in Tragopogon (Roose & Gottlieb, 1976). The three dip- loid species, T. dubius, T. porrifolius, and T. pratensis, were introduced to America from Europe during recent times. They are sharply delimited mor- phologically without overlap in a number of characters (Ownbey, 1950). In southeastern Washington and adjacent Idaho, Ownbey (1950) discovered that interspecific hybridization between them had given rise to two different tet- raploid species: T. dubius and T. porrifolius were the parents of T. mirus, and T. dubius and T. pratensis were the parents of T. miscellus. These two tetraploid species represent the only unambiguous examples of the very recent natural origin of polyploid species. Electrophoretic evidence revealed that the three diploid species in North America are completely divergent (monomorphic for different alleles) at about 40% of the 21 genes that were examined, a result fully concordant with their morphological differentiation. The tetraploids inherited both alleles at each of these genes: T. mirus expresses an additive pattern for nine genes and T. mis- cellus for seven genes, including five in common with T. mirus. In both tet- raploids, the additive gs includes novel hybrid enzymes not produced in the diploid species. Each of the enzyme phenotypes in the tetraploids was fully accounted for by simple additivity of the polypeptides specified by genes in- herited from its respective diploid parents. The observed patterns fully con- firmed the ancestry of both tetraploids which was proposed by Ownbey (1950). These results, as well as others, clearly indicate that electrophoretic analysis of large numbers of enzymes can be an extremely useful and precise probe to iden- tify diploid progenitors of polyploid species. In summary, electrophoretic evidence can be used to test many different types of hypotheses regarding genetic divergence that have already been gen- erated in plant systematics. 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Gene evolution and the hemoglobins, Nature 189: 704. Jacoss, M. 1975. Isozymes and a strategy for their 1 in plant genetics. I. Isozymes: genetic and . 1 ol. Pp. 365-378, in Ledoux (editor), Genetic Manipula- tions with Plant Material. Plenum Press, New Jounson, G. B. 1976. Hidden alleles at je 8 8 glycerophosphate dehydrogenase locus in Colias butterflies. Genetics 83: 149-167 LEVIN, D. 1975. Interspecific hybridization, e and gene exchange in Phlox. Evolution 29: 37-51. 1977] GOTTLIEB—ELECTROPHORESIS AND PLANT SYSTEMATICS 179 X W. L. CnEPET. 1973. Genetic variation in Lycopodium lucidulum, a phylogenetic relic. oe 27: 622-632. Levy, M. & D. A. Levin. 1975. Genic heterozygosity and variation in permanent trans- om . of the Oenothera biennis complex. Genetics 79: 493-512. , E. E. STEINER & D. A. Levin. 1975. Allozyme genetics in permanent translocation heterozygotes n Eu Oenothera biennis complex. Biochem. Genet 0. Lewis, H. & C. ELIN G. 1959. 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Res. 18: 57-69. —0 Nri, M. Genetic distance between populations. pos Naturalist 106: 283-292. —————. 1975. Molecular aera Genetics and Evolution. North-Holland Publ. Co., Amsterdam. Nevo, E., Y. J. Ki, C. R. SHaw & C. S. THAELER. 1974. Pene variation, selection and speciation in Thomomys 1 pocket gophers. Evolution 28: 1-23. OnNpurrF, R. 1976. Speciation and oligogenic differentiation in Lasthenia (Compositae). st. Bot. 1: 91-96. Ownsey, M. 1950. Natural hybridization and amphiploidy in the genus Tragopogon. Amer. J. Bot. 37: 487—499. PowELL, J. 1975. Protein variation in natural populations of animals. Evol. Biol. 8: 9-119. BEC! Press, New Price, S. C. ;enetic vari: ab ility in ecologically marginal and central pu e the 17 elt Clarkia williamsonii (Onagraceae). M.S. thesis, California State Un San Dieg Reppy, M. "E. GARBER. 1971. Genetic studies of variant enzymes. II. Comparative Scc Pa studies of esterases and peroxidases for spedies, hybrids, and amiphinlod: in the genus Nicotiana. Bot. Gaz. (Crawfordsville) 132: 158-166 Rick, C. M. & J. F. Foses. 1975. Allozymes of 5 tomatoes; polymorphism, geo- graphic distribution, and affinities. Evolution 29: —457. E. KEsICKI, F. Foses & M. Horre. 1976. Genetic and 11 studies on | bur new sibling species of Lycopersicon from interandean Perú. Theor. Appl. Genet. 47: 55-68. Rocers, J. S. 1972. Measures of genetic similarity and genetic distance, Univ. Texas Publ. 7213: 145-153. Roose, M. L. & L. D. Gorrrikes. 1976. Genetic and biochemical consequences of poly- ploidy in Tragopogon. Evolution 30: 818-830. ScANDALIOS, J. G. 1969. Genetic control of multiple molecular forms of enzymes in plants: review. “Biochem. Genet. 3: 37-79. 1974. Isozymes in development and differentiation. Annual Rev. Pl. Physiol. ScuaaL, B. A. 1975. 5 structure and local differentiation in Liatris cylindracea. Amer. Naturalist 109: SELANDER, R. K. 1976. Eas vastetian in natural 1 Pp. 21-45, in F. J. Ayala (editor), Molecular Evolution. Sinauer Assoc., Inc., Sunderland, Massachusetts, Suaw, C. R. 1970. How many genes evolve? Biochem. Genet. 4: 275-283 SHEEN, S. 1972. Isozymic evidence bearing on the origin of Nicotiana tabacum. Evolution 26: 143-154 1S0 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Sic, C. F. & G. J. Brewer. 1969. Isozymes of a polyploid series of wheat. Genetics 61: 391-398. SincH, R. S., J. L. Hussy & R. C. Lewontin. 1974. Molecular heterosis for heat-sensitive enzyme alleles. Proc. Ra Acad. U.S.A. 71: 1808-1810. SMrru, H. H., D. E. HAMiLL, E. A. WEAVER & K. H. THOMPSON. 0. Multiple molecular forms of peroxidases and esterases among Nicotiana species d. S J. Heredity 61: TORRES, A. M. 1976. Dissociation-recombination of intergenic sunflower "m dehydro- enase isozymes and relative isozyme activities. Biochem. Genet. 14: 87-9 TURNER, B. 1956. cytotaxonomic study of the genus Hymenopappus TNT Rhodora 58: 164—308. ) Chemosystematics: recent developments. Taxon 18: 134-151. THE APPLICATIONS OF MOLECULAR EVOLUTION TO SYSTEMATICS: RATES, REGULATION, AND THE ROLE OF NATURAL SELECTION' Mary-CLAIRE KING? The development of biochemical methods for comparing the amino acid se- quences of homologous proteins from different species has provided a powerful tool for investigations of evolution and systematics. Perhaps the most intriguing (and most controversial) result of the comparative studies of proteins using these methods has been the discovery that sequences may change at nearly constant rates (Wilson, Carlson & White, 1977). This is not to imply that dif- ferent genes or proteins evolve at the same rate: rather, each class of proteins has its own characteristic rate (Dickerson, 1971). (Serum albumin, for exam- ple, has evolved more rapidly than cytochrome c, but serum albumin has evolved at approximately the same rate among all species of mammals tested, as has cyto- chrome c.) The degree of rate constancy has been the subject of intense debate, but the most current evidence indicates that the variation in evolutionary rate for a given protein is only about twice the variation expected for a totally stochastic process such as radioactive decay (Fitch, 1976). Within these limits, then, a given macromolecular sequence may be used as an evolutionary "clock." The empirical discovery that molecules can be evolutionary "clocks" has been applied to a variety of problems in evolution and systematics. Most frequently, sequence data for a given protein from a number of species has been used to reconstruct phylogenetic trees depicting the probable order of branching of the lineages leading to modern species from a common ancestor. For example, Boul- ter and his colleagues have reconstructed a possible phylogeny for the flowering plants based on the cytochrome c, plastocyanin, and ferredoxin sequences of representative species (Boulter, 1974). In addition, phylogenetic analysis of se- quences of 5S and 16S ribosomal RNA from chloroplasts, bacteria, blue-green algae, and cytoplasm of green plants has confirmed that chloroplasts evolved from photosynthetic prokaryotes living as endosymbionts within the cytoplasms of primitive heterotrophic plants (Margulis, 1970; Bonen & Doolittle, 1976; Zablen et al., 1975; Hori, 1975). Molecular phylogenies may also indicate evolutionary times of divergence if the divergence time for at least one branching event in a tree can be accurately estimated from paleontological or biogeographical evidence. The use of cyto- chrome c sequences to estimate times of divergence for flowering plants poses a fascinating, and still unresolved dilemma. If the cytochromes c of plants are evolving at the same rate as those of vertebrates, which have a unit evolutionary period of about 20 million years, then the intraordinal divergence times for flower- ! I thank A. C. Wilson, S. C. Carlson, and T. J. Tbe for generous contribution of ideas and Du aterik. and A. Hurley for technical assistanc irtment of Biomedical and Environmental Health Sciences, University of California, Berkele s California 94720. ANN. Missourt Bor. Garp. 64: 181-183, 182 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ing plants would be about 240 million years ago (Ramshaw et al., 1972; Boulter et al., 1972). Since the first clearly authentic fossils of flowering plants occur about 130 million years ago (Sporne, 1971), an alternative interpretation of the data is that morphologically plant cytochromes c have evolved twice as fast as those of vertebrates (Cronquist, 1976). Workers in this field are now giving serious attention to the possibility that the origin of flowering plants is more ancient than is indicated by the available fossil evidence (Wilson, Carlson & White, 1977). This situation may be analogous with the origin of mammals, in that the group is very ancient, while adaptive radiation within the group is more recent. The molecular evolutionary approach may also have revealed an important mechanism for evolution at the organismal level. This discovery results from the observed discrepancy between the evolution of macromolecules and the evo- lution of organisms. The comparison of humans and chimpanzees at both the macromolecular level and organismal levels indicates that the two species dif- fer to an extent considered familial in morphology, behavior, and adaptive strat- egy, while their protein sequences differ by less than one percent—a level of difference characteristic of sibling species of Drosophilia or mammals (King & Wilson, 1975). Major adaptive changes may thus be based on molecular events other than sequence changes in structural genes. What sorts of events might these be? Experimental studies of bacterial evolution have demonstrated that major phenotypic changes—in the bacterial case the acquisition of a new meta- bolic activity—depend on an increase in the effective concentration of a protein which previously limited the rate of metabolism of a given substrate, rather than on a qualitative change in the substrate specificity of any protein (Lerner et al., 1964). These quantitative effects could be due to point mutations in regu- latory genes or to chromosomal rearrangements such as duplications and trans- locations (Wilson, 1975). The observation that rates of karyotypic change are fastest in vertebrate groups with the most rapid phenotypic evolution may indi- cate that major adaptive shifts in the evolution of multicellular organisms are frequently associated with chromosomal rearrangements (Wilson et al., 1975). The independence of the evolution of organisms and the evolution of their structural genes may provide a new perspective for investigating the evolutionary roles of natural selection versus random fixiation of selectively neutral alleles. If the random fixation of neutral substitutions were principally responsible for sequence evolution of genes and proteins, it would follow that the rate of se- quence evolution would depend primarily on the mutation rate, which is as- sumed to be constant with time. The “neutral” hypothesis is thus consistent with the observation that sequence evolution depends on calendar time. In addi- tion, the "neutral" hypothesis accounts for the observation that proteins can differ greatly in sequence without differing appreciably in biological activity. At the same time, it is unequivocably established that natural selection acts at the level of the organism, and that this selective pressure varies greatly over time and space. It is tempting to suggest a hypothesis consistent with—though in no way proven by—each of these observations: that fixed substitutions in the vari- able regions of structural genes have generally not been subject to selective 1977] KING—MOLECULAR EVOLUTION AND SYSTEMATICS 183 pressures, since these substitutions are largely irrelevant to the adaptive success of the organism. Instead, natural selection at the level of the organism may be reflected at the molecular level in both rapid elimination of deleterious muta- tions in regulatory systems and fixation of occasional adaptive changes in loci or patterns of genome organization controlling the expression of structural genes. LITERATURE CITED Bonen, L. DoorrrrLEe. 1976. Partial seque p of 168 rRNA and the phylogeny of ei Ec and chloroplasts. Nature 261: 669-673. BovrrEn, D. . The t de of plant proteins qi Special reference to higher plant cytochrome c. Curr. 2 vances Pl. Sci. Comment. Pl. S : 1-16 AMSH E W. 'THOMPSON, M. ae & R. H. Brown. 1972. A phylogeny of higher 7 based on the amino acid sequences of cytochrome c and its biological i mplications. Proc. Roy. Soc. London, Ser. B, Biol. Sci. 181: 441—455. 976. The taxonomic significance of the structure of plant proteins: a Mie. classical taxonomist's view. Brittonia 28: 127. Dic gen .E. 19 The structure of cytochrome c and the rates of molecular evolution. c. Evol. 1: 5. FrrcH, Ex "M. 1976. The molecular evolution of cytochrome c in eukaryotes. J. Molec. Evol. 8: 13-40. Hee H. 1975. P e “i i RNA. J. Molec. Evol. 7: 75-86. KIN = C. & A. C. Wir 975. Evolution at bo ia. in humans and chimpanzees. nce 17 107-116 "Sci Lenen, 3 A., . Wu & E. C. C. LIN. 1964. Evolution of a catabolic pathway in bac- Scene 146: 1313-1315. E sawsi L. 1970. Origin of Eukaryotic Cells. Yale Univ. i New Haven. RANISHAW, J. A. M., D. L. RicHARDsoN, B. T. MEATYARD, OWN, M. RICHARDSON, E. W. THoMPsov & D. BOULTER. 12. ‘The tive of — of the ‘flowering plants de- m by using amino acid sequence data of cytochrome c. New Phytol. 71: 773— 779. 1. rc WILSON, A. C. 1975. Evolutionary importance of gene regulation. Stadler Genet. Symp. i 9 77: 117-134. Univ. of Missouri, Columbia. , S. S. CaRLSON & T. J. Wurrr. 1977. Biochemical evolution. Annual Rev. Biochem. MUR K. R. 1971. The Mysterious Origin of Flowering Plants. Oxford Univ. Press, Ox- for M. Cas M. C. KiN 1975. Social structuring of mammalian Proc. Natl. Acad. U.S.A. 72: 5061-5065. ZABLEN, L. f. S. KissiL, C. R. Worse & D. E. Burrow. 1975. Phylogenetic origin of the e and prokaryotic nature of its ribosomal RNA. Proc. Natl. Acad. U.S.A. 72: 2418-2422. Busnu, S. 8 and rate of Bs scias: ut B., N CHEMOSYSTEMATICS—ANALYSES OF POPULATIONAL DIFFERENTIATION AND VARIABILITY OF ANCESTRAL AND RECENT POPULATIONS OF JUNIPERUS ASHEI' RoBERT P. ADAMS” ABSTRACT Three types of data were used to analyze 28 natural populations of Juniperus ashei: 16 morphological characters, 152 terpenoids, and 23 peroxidases. In this paper the peroxidase electromorphs were treated as ordinary qualitative chemical characters to examine the feasi- bility of using isozymes for taxonomic purposes and as indicators of pop dal variability. a sets were subjected to various numerical analyses to examine regional trends, an- inities i it po ) e mi f i Coordinate loadings were then contoured for the first three coordinates of each similarity matrix to aid the visualization of the regional trends. The terpenoids pis UD showed a series of uniform populations from central Texas into the Ozarks icu ee popula- tions on the south and west portions of the range, extending into no 1 5 exic o sip trends were apparent in the peroxidases and no corresponding modes of variation De ta i ; A W are discussed. ee 8 body 0 n ashei 5 from central Texas to the Ozarks appear to be A (recent), hse the divergent populations seem to be more primitive (ancestral). A method called differential similarities is introduced to analyze the clinal grada- tion of J. ashei toward J. saltillensis in Mexico. Intrapopulational variability was analyzed In general, the rec nt populations had high 1 8 and low variability, and the cal and terpenoid du. The pattern of variation in the ordo could not be generalized ut ran 0 n oxid upon, but a to be mosaic. Per ses < not appear useful in this analysis when subjected te mn kosy pey 5 he evolution of J. ashei into its present distribution appear. least tv ie composed of very uniform, recent migra- tions and pers usns inen velie population perhaps extending close to the geographic origin of this taxon in northern. Mex The use of chemical characters has gained widespread acceptance during the past decade to the point that a graduate student thesis in systematics is now un- usual if no chemical data are utilized. Because of the relative ease of use, flavo- noids are widely utilized in systematic and evolutionary plant studies. The early works on Asplenium (Smith & Levin, 1963), Lemnaceae (McClure & Alston, 1966) and Baptisia (Alston & Turner, 1963) are classics, required read- ing for chemosystematic students. Likewise, classic is the work on betalains by Mabry and coworkers (summarized in this symposium). Whereas flavonoids and betalains have been extensively used above the species level (probably due to the qualitative nature of the methods), terpenoids, due to the quantitative nature of gas/liquid chromatography, have been more widely used at or below thank Walter Kelley for field assistance and the use of the peroxidase data which he diligently gathered. This research was supported by NSF Grants GB24320 and GB37315X. C omputer time was furnished by Colorado State University. Science Research Center, Hardin-Simmons University, Box 1095, Gruver, Texas 79040. ANN. Missouni Bor. Garp. 64: 184-209. 1977. 1977] ADAMS—JUNIPERUS ASHEI 185 the species level. The gymnosperms have been the focus of many studies of populational differentiation which have uncovered clines (Flake et al., 1969, 1973), chemical races (Smith et al., 1969), hybridization (see von Rudloff, 1975 for an excellent review), and ancestral migrations (Adams, 1975a; Zavarin & Snajberk, 1973). In the angiosperms, work on the monterpenes of Bursera (Mooney & Emboden, 1968) demonstrated the use of these compounds in the detection of clinal variation. Of course, the work on the Australian Eucalyptus species has been of tremendous use in the classification of populations of these taxa and is well known. Although studies using terpenoid characters to analyze populational differ- entiation are well known, the analvsis of population variability in relation to im- portant population biology questions such as the founder's effect, genetic drift, the effects of small versus large populations and central versus peripheral sites on variability have not been addressed. The relatively recent rise in the use of isoenzyme data has rekindled an interest in the examination of these questions. Gottlieb (at this symposium) has reviewed the literature on isozymes and their use in 5 Nevertheless, it seems in order to mention that the “isozyme bandwagon” has become the current fad before we have developed à very thorough knowledge about the molecular basis of the electromorphs distinguished on gels, Before the widespread use of isozymes, the study of variability within popu- lations seems to have stagnated with the exception of the numerical taxonomic school (including morphometrics). Gilmartin (1969a, 1969b, 1974, 1976) has introduced a new idea called the coefficient of phentic variation (CPV) to ex- amine the combined effects of many characters on variability. The CPV is merely the standard deviation of the mean similarity among a group of opera- tional taxonomic units (OTUs) divided by that mean similarity. Whereas the mean similarity of a group tells about the average affinities, the CPV shows how homogeneous are the similarities of one group versus another group. Since the CPV is normalized by the mean similarity, different character sets can be com- pared as well as different levels of organization (i.e., population vs. species vs. nus). To my knowledge, the CPVs have not been used to study population vari- ability with the exception of the studies by Gilmartin. The purpose of this paper is to examine population differentiation and variability in Juniperus ashei Buch. using three contrasting sets of characters: morphological characters, volatile terpenoids from leaves, and leaf peroxidases. The literature on J. ashei has been reviewed by Adams & Turner (1970 Juniperus ashei is a taxon of a rather restricted range, occurring on lime- stone outcrops from northern Mexico to southern Missouri (Fig. 1). The Ed- wards Plateau region of central Texas supports dense populations covering thousands of acres, whereas the disjunct populations ( Lubbock-Post, Texarkana, Arbuckle Mountains, Ozark Mountains, and northern Mexico) often have nearly pure stands of J. ashei, but seldom cover such large areas. Being a fairly con- spicuous conifer tree, one can be relatively confident in the taxonomic distribu- tion records which imply that there are few, if any, trees between the disjunct populations and the Edwards Plateau populations. Thus, this would appear to 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 MISSOURI 2 N A ay AP WY DA OKL AHOMA ME AD w 2222 | MIN SAS N 28 SSS ®OKLAHOMA CITY 4 f ARBUCKLE MTNS. 227 TEXAS ~}TEXARKANA I#-UBBOCK eDALLAS "954 ABILENE ARKANSAS KILOMETERS SIERRA DEL CARMEN MTNS. p Ficure 1. Distribution of Juniperus ashei showing the 28 populations sampled for this study. The exact oun on of J. ashei in northern Mexico is not known and is indicated generally by a dashed line be an excellent taxon to test some of the hypotheses advanced by Ehrlich & Raven (1969) in regard to gene flow versus selection in the maintenance of species. Previous research (Adams & Turner, 1970; Adams, 1975a) has shown that the terpenoids of this taxon exhibit a remarkably high similarity between central Texas and the Ozarks (Fig. 2). However, many significant differences were found between populations 12, 13, and 17 and the other populations. One tree of J. ashei (number 116 in Fig. 2) was discovered in Mexico and found to cluster with the atypical populations (12, 13, 17). This, along with similar evidence in J. pinchotti populations (Adams, 1975b) seemed to imply that relicit migrations have been very important in the establishment of these patterns. Evidence from rat middens and palynology in the southwestern United States is considerable (King, 1973; Mehringer et al., 1970; Van Devender & King, 1971; Wells, 1965, 1966, 1970; Wells & Berger, 1967; Whitehead, 1972; Wright, 1970) 1977] ADAMS—JUNIPERUS ASHEI 187 MISSOURI ee T — OKLAHOMA Z = A. ifa Let I TEXAS ARKANSAS MONONA Oo O D p —— NEW Z — ^A BRAUNFELS .Z = 7 — Z Z Q 9 2 Z TA A ta / FIGURE 2. Contoured similarities based on 54 terpenoid characters, F-1 weighted. Notice the uniformity from central Texas to the Ozarks and the clustering of Dunal mos 12, 13 and 17 with tree 116 from northern Mexico (from Adams, 1975a). that the Pleistocene ice advances pushed boreal and temperate species to lower elevations and southward. The northern Chihuahuan desert was certainly in- vaded by Juniperus (Wells, 1966) and crossed repeatedly. Even so, the data presented by Adams (1975a) for the close similarities of populations 12, 13 and 17 to northern Mexico J. ashei have remained somewhat tentative. This is due to the use of only 1 tree (number 116) from northern Mexico and the fact that no morphological data were used except in the largely preliminary study by Adams & Turner (1970). In this study I will remedy these shortcomings by reporting on 15 trees of J. ashei from northern Mexico (population 25 in Fig. 1), as well as 4 additional populations: Post (near Lubbock), 24; Pandale, 26; Texarkana, 27; Saline Creek, Oklahoma, 28 (see Fig. 1). In addition, I report data on 16 morphological 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 LE Sixteen morphological characters and states scored for 15 trees from each of the 28 populations of J. ashei sampled (Fig. 1). Missing data was coded by a -1.0 for a flag in statistical analysis. Character States (if applicable) FDI FEMALE CONE DIAMETER: avg. of up to 10 and not less than 4 (in mm). FCO FEMALE CONE COLOR: 1.0-4.0 (blue-yellow/brown ). SPF SEEDS PER FEMALE CONE: avg. of up to 10 cones scored, not less than 4. BLM BLOOM ON CONE: 1.0-3.0 (none to very heavy coating). SEA SEED AREA: seed length x width, avg. of 10 seeds and not Vu than 4. SER SEED WIDTH/LENGTH: avg. of 10 seeds and not less than WGA WHIP LEAF GLAND AREA: whip leaf gland width x E avg. of 5 glands. WGR WHIP LEAF GLAND LENGTH/WIDTH: ratio, avg. of 5 glands. WLM WHIP LEAF MARGINS: 1.0-4.0 (smooth-heavy serration) avg. of 5 les vas. WGP WHIP GLANDS/PROTRUSION: 1.0-3.0 (sunken-smooth-protrudes), avg. of 5 glands. WRP iiec n RUPTURED: 1.0-3.0 (none-some-almost all), avg. of 5 ob- ervation B/S WHIP LEAF BLADE LENGTH/SHEATH LENGTH: avg. of 5 leaves. G/S WHIP LEAF GLAND pedir iu LENGTH: avg. of 5 leaves. SLL SCALE LEAF LENGTH: avg. of 5 lea L/B SCALE LEAF LENGTH/BRANCH WID DTH: Ratio of scale leaf length to the width of the branch (twig) where that scale leaf was borne. Avg. of 5 mea- surements. BAN BRANCHING ANGLE: Angle of branching of ultimate twig, avg. of 5 measure- ments (each to nearest 5 degrees). characters, as well as peroxidases, from leaves. Finally, I compare these 3 sets of characters both in regard to their use in the analysis of populational differentia- tion and in the analysis of variability within populations. MATERIALS AND METHODS Twenty-eight populations of J. ashei were sampled throughout the natural range (Fig. 1). For the terpenoid and morphological characters, 15 trees were sampled from populations 1 through 23 in December, 1970, and 15 trees were sampled from populations 24 through 28 in December and January, 1974-1975, to complete the sampling. The sampling methods are given in Adams & Turner (1970), except that in 1974-1975, the foliage was generally frozen within a few hours in the freezer of our field trailer. Voucher specimens are on file at Colo- rado State University. All samples from each of the two sampling periods were placed in a random sequence for distillation as advocated by Adams (1975c). These procedures convert the temporal changes in foliage, oils, columns, etc. to random variables. Therefore population differentiation patterns can be readily separated from experimental procedural errors in the statistical analysis phase. The volatile terpenoids were steam distilled for 2 h as outlined by Adams (1970) and the extracts were kept at —20°C until analyzed by gas/liquid chromatography. Separation was made on a 200 ft x 0.02 in. capillary column (wall coated with PEG 20M ) as described by Adams (1975a). The identities of the terpenoids of J. ashei are given in von Rudloff (1968) and Adams & Turner (1970). In- dividual peaks were quantified with an electronic digital integrator and auto- matically punched onto computer cards. 1977] ADAMS-—JUNIPERUS ASHEI 189 Sixteen morphological characters were scored as outlined in Table 1 for 15 specimens of 28 populations. Some fruit (female cones) and seed characters were not scored (and were thus set to —1.0 as a flag) since not all trees sampled had female cones. One hundred and forty-two terpenoids were subjected to analysis of variance (ANOVA) to determine which characters showed significant differences among populations. Fifty-nine terpenoids had F ratios greater than 1.0, a maximum population average greater than 0.1% and were used to compute F-1 weighted (Adams, 1975c) mean character differences (MCD or Manhattan metric) simi- larity measures between populations (see Adams, 1972, for exact formulation ). This similarity matrix (28 x 28) was then used as inpnt for principal coordinate analysis ( Gower, 1966, 1967; Williams et al., 1971) to factor the similarity matrix into major coordinates of variation. The first 3 principal coordinates were used to contour map populations as they were ordinated on each of the orthogonal axes. The 16 morphological characters were also analyzed by ANOVA and the Student-Newman-Keuls (SNK) multiple range test was applied (P = 0.05) to determine which populations were significantly different. Fifteen morphological characters (FEMALE CONE COLOR was omitted, F = 0.88) were used to compute a similarity matrix which was then factored by principal coordinates. The first 3 coordinates were contour mapped as outlined above. For the peroxidase work, foliage of 30 plants (occasionally less, see Kelley 1976) were sampled from 15 populations in November-December, 1974, and frozen in the field trailer within a few hours. This foliage was kept frozen until extracted. The enzymes were extracted by grinding the foliage in liquid nitrogen with alumina then adding an extraction buffer of 0.10 M trismaleate, pH 7.00 containing: 0.02 M sodium tetraborate; 0.25 M sodium ascorbate; 0.02 M sodium meta-bisulfite; 0.02 M sodium diethldithiocarbamate (DIECA); 0.01 M ger- manium dioxide; 10% (v/v) dimethyl sulfoxide (DMSO) plus polyvinylpoly- pyrrolidone ( PVPP), 10 gms/50 ml buffer. The complete instructions are lengthy and the interested reader is referred to Kelley (1976) and Kelley & Adams (1977a) for complete details. The peroxidases were concentrated and electro- phoresed on acrylamide gels (discontinuous 4.5, 6, and 8% anodic, see Kelley, 1976) within 72 hours from the time of extraction. Although Kelley (1976) ana- lyzed peroxidases, esterases, and an alcohol dehydrogenase, I am only using the peroxidase data since it showed much of the same pattern of variability as the other systems (Kelley, 1976). Peroxidases in Juniperus are little effected by sea- sonal differences (Kelley & Adams 1977a), and peroxidases are generally very stable (Kelley, 1976). Peroxidases were stained with o-tolidine/ HzO: (Denna & Alexander, 1975). An aggregate total of 23 peroxidase bands were found in the 15 populations of J. ashei sampled. In cases where bands were very close together on the gel, samples were corun to determine which electromorphs were different. These bands were each scored as 1.0 (present) or 0.0 (absent) for each plant and then subjected to ANOVA to obtain some estimate of F ratios for character weighting. Of course, ANOVA of qualitative data has a tendency to underestimate the F ratios, but this did provide a crude method to obtain relative character weights. Sixteen peroxidases had F greater than 1.0 and were ANNALS OF THE MISSOURI BOTANICAL GARDEN 190 @ MISSOURI OKLAHOMA 28e| 3[*2| 1 7 RKANSAS *4 ; 222 ePOST 5 XARKANA 7 H je 2 1 Q e7 TEXAS 7 2 8 ZS SUM 19 198 , ° 18e 9 e e 13 15 189] 0. Q 199 290 5e KILOMETER. COO (SO /), TERPENES 4 5 5 5 6 7 š; (5) MISSÓURI 2 OKLAHOMA 2800 | 31% 1 d 4 ISAS , 7 T ° e2 4 TEXARKANA Ñ 1 24e 05 2 e 3 Ne © 3 4 o ^ Oe e7 ma e $ *8 TEXAS 4 26e U 18 e9 1 " . 5 25 4 [6 17 Q 199 290 ° MEX. KILOMETERS ° > 7) WussSuRi OKLA A 6 280 > 1 A NSAS e4 I 7 m 7 TEXARKANA 24e e^ ° a KA 5 2 4 D e7 3 e 1124 Ca TEXAS 26.8 Q Q V° 2 4 2 e /e ° Š 115) 16 7 O 100 200 $ MEX NLOMETERS COORD .2(994) TERPENES | COORD 3($94), TERPENES [Vor. 64 6 7 @ fuso OKLAHOMA e 3 2 1 5 ARKANSAS 04 e27 NU 7 240 š H 22 2 q ° e7 TEXAS Yes. ° 4 ; 8e ; 15 je 5 Q "o 290 5e KILOMETERS $ COORD.1 (389/9)), MORPH. 4 5 6 3 6 MISSOURI 7 AHOMA 2 2211 4 KANSAS 5 e4 : 2 . TEXARKANA! 1 240 5 1 3 e 2 ‘ 5 (° " oe 93 2 U A19 TEXAS Ë 4 2 ° ge e. 4 3NA 15 16 17 © 100 200 A KILOMETERS X, i } 5 COORD. 2(9°%o), MORPH. 2 2 3 4 5 2 mI OKL A 1 AS e4 6 XARKANA ‘V7 1 240 e5 P 21 : ; 3 4 > Oe š p . /e. PO Texas 6 08 1 745 ° 01 16 O 100 200 25 — — ° KILOMETERS MEK. 4 4 4 COORD. 3(8 %o), MORPH. 1977] ADAMS—JUNIPERUS ASHEI 191 not uniformly unique to one population. Similarity measures were computed as outlined above, and the similarity matrix (15 X 15) was factored to obtain principal coordinates. The first 2 coordinates were contour mapped for com- parison of regional trends. For the analysis of within populational variability, 3 sets of similarity measures were calculated using all terpenoid, morphological, and peroxidase characters, equally weighted. It appears that F weighting is not desirable when examining intrapopulational variation. These analyses resulted in 3 kinds of similarity matrices (terpenoid, morphological, and peroxidase) for each population. The average similarity (Sr) was then computed for each population along with the coefficient of phenetic variation (CPV = Sd—/Sr ). The Sr's and CPVs were then contour mapped to examine regional trends of intrapopulational variation. POPULATIONAL DIFFERENTIATION The principal trend in the terpenoid similarities is that of the differentiation of populations 25, 26, 12, 13 and 17 from the rest of the populations (Fig. 3). From these coordinate loadings one can see (Table 2) that 50% of the variation in the similarities is mostly due to the divergent nature of populations 25, 26, 12 13 and 17. The high negative loading of population 17 onto coordinate one in- dicates that population 17 (New Braunfels, Texas) has considerable affinities with the west Texas and Mexico plants. It is interesting to compare the major trend of the terpenoids with that of the morphology (Fig. 3 vs. Fig. 4). This major trend in the morphology accounts for 38% of the variation in similarities and is practically identical to the major trend of the terpenoids. A couple of ex- ceptions are that the Post population (24) seems more similar to the west Texas- Mexico populations in the morphology, while the New Braunfels population (17) is not quite as different from the central Texas populations in its morphology as in its terpenoids. In both cases, from central Texas to the Ozarks a picture of uniformity is presented. It might be noted that this compares very closely with the contoured terpenoid phenogram Fig. 2 (from Adams, 1975a). It appears that the major coordinate of principal coordinates analysis is the dominant — 1 — < FicurEs 3-8.—3-4. Contoured loadings of S wa l coordinate 1 extracted from similar- ity measures among populations, (see Tables 2-3).—3. This pattern extracted 50 0% of the variation from the terpenoid similarity matrix. s n this pattern is the Principal pattern previously shown (Fig. 2). Contours: 1 = -0.70; 7 = 0.18.—4. This pattern accounted for 38% of the variation from the ee S. similarity matrix. The Post sal. ation (24) shares some affinities to the west Texas-Mexico populations. Populatian 17 seems a S less divergent in its morphology than its terpenoide | Fig. 3). Contours: 1 = —0.64; 7 13.— 5-6. Es red principal coordinate 5 ius trend (9%, terpenoids) seems e "n yo to 85 ` divergence of populations 24, 27, and 3 8, plus sampling differences (see text). Contours: -0.37; 7 = 0.16.—6. This t rend (9%, morphological) seems to be due | to procedural road in scoring the Bose rient characters (see text). Contours: 1 = -0.32; 7 = .16.—7-8. Contoured principal n ate 3.—7. Divergence of the Post (24) population is most evident A edge this coordinate (5%, terpenoids). Contours: 1 = -0.2 0.28.— Note the str mg e nce EL populations 27 and 28 (8%, -— l Con- — 1 = -0.22; 0.2 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE 2. Principal coordinate analysis of similarity matrices using terpenoids, morphol- ogy, and peroxidase characters for the computations of similarity measures between popula- tions of J. ashei. All eigenroots were extracted from each matrix until they failed to converge. It is thought that when eigenroots begin to level off in values, additional roots represent only random error variance. TeRPENOIDS (Srs based on 59 terpenoids, 28 x 28 matrix) 7196 of variation extracted by 5 roots. Eigenroots 3.56 0.66 0.36 0.29 0.25 variation extracted 496 92 51 40 3.6 MonPnoLocy (Srs based on 15 morphological characters, 28 X 28 matrix) 72% of variation removed by 7 roots. Eigenroots 2.40 0.64 0.53 0.36 0.34 0.30 0.26 % variation extracted 35.9 95 8.0 ; PEnoxipAsES (Srs based on 16 peroxidases, 15 X 15 matrix) 93% of variation extracted by 10 roots. Eigenroots 1.68 085 0.62 0.53 0.35 0.30 0.27 0.21 0.19 0.18 % variation extracted 301 152 112 95 63 54 48 . 35 3.2 theme of a single linkage phenogram (see Adams, 1975a). Thus, we see that the west Texas-Mexico type populations account for 50% and 38% of the variation in the terpenoid and morphological similarities, respectively. The second coordinate extracted from the terpenoid similarity matrix largely separates the small island populations at Post (24), Texarkana (27), and Saline Creek (28) from the rest of J. ashei (Fig. 5). These populations, along with 25 and 26, were collected and analyzed 4 years later (1974) than the other popu- lations (1970 collections). Therefore part of these differences may be due to sampling methods, seasonal variations, and different gas chromatographic con- ditions. However, populations 25 and 26 seem to cluster well with populations sampled in 1970, so this factor may be only a minor cause of this trend. It seems that this small amount of variation (9%) is chiefly accounted for by the diver- gence of these 3 small, isolated populations (24, 27, 28), along with a contribution resulting from different sampling and analysis times. The second coordinate of the morphological similarity matrix (Fig. 6) is clearly due to the fact that popu- lations 24, 26, 26, 27, and 28 were sampled and analyzed in 1974 rather than with the other populations (sampled and analyzed in 1970). It is felt that most of these differences (approximately 9% of the variation in the similarity matrix) are due to the fact that a different technician measured the morphological characters of populations 24, 25, 26, 27, and 28 (1975) than the other populations (1970- 1972). Even with close supervision and training, it is very difficult to get two people to score morphological characters in the same manner. My experience has been that comparisons between morphological data sets scored by com- pletely different research projects is almost impossible. If we consider that the eigenroots of about 5% may be mostly random noise (see below), then the 9% of coordinate 2 is only about twice the experimental error but 25% the size of the major trend. The third coordinate does not appear to be very significant in the terpenoid 1977 ADAMS--JUNIPERUS ASHEI 193 similarity matrix since only 5.1% of the variation was extracted and the eigenroots have leveled off at this value ( Table 2). Contouring of this coordinate ( Fig. 7) shows that most of the variation along this axis is due to population 24 at Post. This population is on one of the most unusual sites that I have seen for J. ashei. It is in a deep ravine, cut into the Permian red clay, just east of the Llano Estacado. The stand is occasionally mixed with J. pinchotii, with J. ashei found in the more mesic spots. This trend could represent a response to microhabitat selection or environmentally induced plasticity. Transplant studies will probably be needed to answer this question. Another trend is that the northern-most (including Post) populations seem to be more heavily loaded onto this coordinate than those populations in the central and southwestern portion of the range. The third coordinate of the morphological similarity matrix extracted 8.0% of the variation and might be significant as the 4th through 7th roots seem to have asymptoted to about 4 or 5%. The contour map of this coordinate (Fig. 8) shows a northwest-southeast trend across the populations, somewhat like that in Fig. 7, except there is a decided split between the Texarkana population (27) and those to the north and west. This population (27) is almost as atypical for J. ashei as the one at Post, Texas (24). At population 27, J. ashei is found on a small ( few acres?) limestone outcrop that is gently sloping and very moist ( 1,143— 1,270 mm of precipitation per year). It is a mixed stand with some J. virginiana. Whether this pattern represents some small microhabitat selections or environ- mentally induced plasticity in the morphology must await transplant studies for additional information. In any case, it is obvious that the major trend in both the terpenoids and morphology is the differentiation of populations 25, 26, 13, 12, and 17 from the rest of the species. Principal coordinate analysis of the similarity matrix based on peroxidases (Tables 2-3) yielded quite different results. A most notable difference being that 10 eigenroots were extracted from a 15 X 15 matrix, whereas only 5 and 7 roots accounted for most of the definable variation in the much larger (28 x 28) matrices of the terpenoids and morphology. This seems to indicate that the peroxidases are varying in many different directions, whereas the terpenoids and morphology seem to display much more directional or concurrent variation. Another interesting facet is that the eigenroots of the terpenoid and morphological similarity matrices quickly decreased to rather constant values after 2 and 3 roots, whereas the roots of the peroxidases seem to tail out much farther. This seems to imply a considerable amount of independence among the peroxidases. Exami— nation of the first coordinate of the peroxidase similarity matrix (Fig. 9) reveals a northeast-southwest pattern. (remember that only the 15 populations marked with an asterisk were analyzed for peroxidases). The Texarkana (27) and Junc- tion (10) populations are most similar to each other, and the Ozark populations (1, 2) are most similar to the north Texas (5, 7) and west Texas-Mexico popula- tions (12, 25). This trend is unlike any other seen in either the morphological or terpenoid data. The divergence of the Texarkana population (27) from the Ozark (1, 2) and north Texas (5, 7) populations would be easy to explain (if one ignores the morphological and terpenoid data) as genetic drift and/or 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ABLE 3. Coordinate loadings (principal coordinates 1, 2, 3 in each case) for popula- tions onto coordinates. These coordinate loadings were used for generating the contour maps in successive figures. Note the close correspondence between coordinate 1 for the terpenoids and morphology. Values in parenthesis indicate the amount of the variation in the Sr matrix accounted for by each of the coordinates. Terpenoids Morphology Peroxidases C; C: C; C; C: C; C; C: Cs Population (50%) (9%) (5%) (38%) (9%) (8%) (3096) (1596) (11%) 1 011 0.18 0.13 016 0.06 0.15 052 0.06 0.11 2 -0.02 0.20 0.08 013 0.13 0.10 0.32 —0.17 3 017 0.12 0.03 0.06 021 -0.15 — — — 4 0.15 0.11 0.04 0.13 0.06 -0.02 -0.16 0.21 -0.14 5 013 O17 0.02 0.12 0.00 0.00 0.44 0.06 0.12 6 0.23 -0.08 -0.09 0.18 0.09 0.01 — — — 7 013 0.18 0.10 0.20 -0.02 0.11 0.38 -0.04 0.02 8 0.15 0.04 -0.14 0.20 0.09 0.15 — — — 9 0.11 0.18 0.01 0.17 -0.05 0.11 -0.27 -0.51 -0.05 10 0.21 -0.04 -0.14 013 0.03 0.10 -0.50 0.20 -0.20 11 011 0.15 0.06 -0.4 0.27 -0.14 — — — 12 -0.84 -0.05 -0.03 -0.71 028 0.20 0.31 0.16 -0.04 15 -0.78 -0.03 -0.03 -0.49 0.18 -0.02 -0.06 -0.06 0.04 14 021 -0.14 -0.20 0.20 0.06 0.04 — — — 15 0.20 -0.05 -0.10 0.26 -0.04 0.10 — — — 16 0.18 0.06 0.02 0.18 -0.05 —0.04 17 -0.67 0.02 —0.00 0.23 0.05 —0.06 -0.33 -0.34 0.10 18 018 0.03 —0.08 0.14 -0.05 0 — 19 0.10 0.16 —0.03 0.09 0.00 -0.07 — — — 20 0.22 -0.23 -0.18 0.17 -0.05 0.03 — — — 2] 026 -0.21 -0.13 0.17 0.01 -0.15 -0.02 -0.11 -0.28 22 0.19 0.07 —0.07 0.13 —0.07 —0.18 — — — 23 0.18 0.08 0.07 0.14 0.03 —0.17 — — — 24 0.22 —0.42 0.32 -0.14 -0.34 —0.27 -0.11 0.30 -0.42 25 -0.78 -0.07 —0.07 -0.71 -0.16 -0.06 018 0.17 0.15 26 -0.71 -0.05 0.08 -0.72 -0.26 0.13 — — — 27 0.20 -0.26 0.11 0.16 -0.36 0.31 -0.58 0.34 0.49 28 0.16 -0.13 0.19 0.02 —0.09 -0.26 0.14 -0.28 0.07 founder’s effect, but the peroxidase similarity to the Junction population (10) rather stretches the point. Coordinate two of the peroxidase similarity matrix shows (Fig. 10) high loadings of populations 27, 4, 24, 10, and 25. This coordinate seems to be a ran- dom assortment of populations distributed across the range of J. ashei. Similar variation (high similarities across disjunct populations and a random mosaic pat- tern) has been previously observed in nonsignificant variation of individual morphological characters (see Adams & Turner, 1970, for several contoured morphological characters). Coordinate three shows another pattern of mosaic variation and the interested reader is referred to Kelley & Adams (1977b) for more detailed maps of peroxidases, esterases, and alcohol dehydrogenases. low can we interpret these conflicting results? One way to view geographical variation is to consider the number of gene differences needed to produce the observed changes. For morphological characters, Charles & Goodwin (1943) have shown that in Solidago many morphological characters used in taxonomy 1977] ADAMS—JUNIPERUS ASHEI 195 4 5 6 7 5 4 3 3 4 6 (9) MISEOURI (9 SSOURI OKLAHOMA O. A z i ane ie 1 28e e2 s TEXAS n TEXAS ANJAS 6 5 5 L^! 5 e4 6 š 6 7 H post ARKAN POST TEXARKANA |7 5 1 ° , 21 24 Ç 2 e^ e6 < ° 6 76 e7 6 C 4 23 7 1 Me 2661 19 e . T %9 $ > 6 ia SS 5 1 ©17 3 6 e, n — 299 4 25 KILOMETERS 23 EX —— N PEROXIDASES COORD. 289%) PEROXIDASES Ficures 9-10.—9. kind ie as loadings of principal coordinate 1, extracted from the F-1 weighted peroxidase similarity measures among populations (see Tables 2-3). This co- ordinate extracted 30% of the à deoa from the matrix. Only those 15 populations pu with an asterisk were analyzed for the peroxidases. See text for VP du Contours: = -0.51; 7 = 0.43.—10. Contoured principal 5 2 (15% of the variation, ns Saatis), No regional trends were uncoverec this or any of the successive coordinates extracted. See text for discussion. Contours: 1 = 0.42 7 = 0.2 are controlled by a minimum of 4, 5, and 6 genes. Irving & Adams (1973) in a study of Hedeoma terpenoids found that those terpenoids were controlled by a minimum of 1, 2, and 3, but up to 7, genes which agrees with the work on Pinus by Hanover (1966) and others. The peroxidase electromorphs isolated on gels represent probably no more than 1 gene for each 2 bands in the composite. Sup- pose we assume that the 15 morphological characters are each controlled on the average by 5 genes, the 59 terpenoid characters each are controlled by 2 genes (average), and the 16 peroxidase bands are each controlled by 1 independent allele, with 2 alleles (simple codominance) per gene. This means that the pat- tern displayed by the morphological data sampled a minimum of 75 genes, with a minimum sample of 118 genes for the terpenoids, and a maximum sample of 8 genes for the peroxidases. Of course, we have ignored pleiotropy, epistatis, and linkage, but we have no a priori knowledge that these factors are of differential genetic importance in any of these 3 kinds of data. To obtain a random sample of the genome, one would have to favor the morphological and terpenoid data on the basis of sample size alone. Together the morphology and terpenoids (minimum of 193 genes) overshadow the peroxidase data (maximum of 8 genes). Even so it is striking that no logical regional trends emerged from the peroxidases (nor from the esterases or alcohol dehydrogenases, Kelley & Adams, 1977b). The problems of homology may account for much of this random similarity be- tween widely, disjunct populations (e.g., population 10 and 27, Fig. 9). Ho- mology between the morphology of these populations (Table 1) is practically assured. The terpenoid variation is almost totally quantitative in this taxon, and 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 the resolution obtained with capillary gas chromatography greatly increases the probability that peaks from different populations of a quantitatively varying species are in fact the same compound (although there is a small finite prob- ability that different genes produce the same compound in different populations). On the other hand, the peroxidases were often found to be qualitatively varying between close, adjacent populations with a band being in very high frequencies in one population and totally missing from our sample in another population. The high similarities obtained in mosaic patterns (Figs. 9-10) are most readily explained by lack of homology between peroxidase bands, although parallel microselection could play an important role. As far as I know, there have been no cases showing that electrophoretic mobility, per se, is under selection (that is not to say that proteins bearing more positive or negative charges might not be selected due to substrate affinity, etc.). Four hypotheses have been advanced (Adams & Turner, 1970) to explain the pattern of regional variation seen in the terpenoids and morphology of J. ashei. Two of these, sampling errors and parallel selection (in populations 17, 12, 13, 25, 26) have been pretty well disposed of by Adams (1975a). The other two, predominately southerly winds during pollination (December-January ) and northward bird migration during the spring, and ancestral migration leav- ing relict populations, deserve additional discussion. The prevailing wind dur- ing the pollination period ( December-January) is generally from the south on the Edwards Plateau (Arbingast et al, 1967). Thus, one might expect pollen to be generally blown northward. Coupled with the northward migration of Cedar waxwings and other birds that feed on J. ashei berries (female cones), this would tend to isolate population 17 from breeding with adjacent popula- tions (9, 16, 18). This would also help explain the north-south line of differenti- ation between populations 12, 13 and 11, 14 (Figs. 3-4). Although these phe- nomena help explain the persistence of the pattern, they do little to explain the common patterns seen in populations 25, 26, 12, 13, and 17. Ancestral migrations leaving relict populations could help explain these patterns. PLEISTOCENE PATTERNS Although there is considerable evidence of a continuous band of sclerophyl- lous vegetation from central Texas into northern Mexico during the Tertiary (Axelrod, 1975), I would like to focus on events of the Pleistocene, particularly the last pluvial and interglacial periods. I have reconstructed parts of the vege- tation during the Wisconsin pluvial, 10,000-20,000 B.P., in Fig. 11. According to King (1973) the western Missouri Ozarks were covered with boreal spruce forest from about 25,000 to at least 13,000 B.P., with pine parkland preceding the boreal spruce. Since the pine parkland and boreal spruce forest both ap- pear to have been pushed southward from the north (Dillon, 1956), I have as- sumed that the area south of the Ozarks may have been pine woodland or park- land (also see Bryant, 1969). A pine-spruce woodland seems likely in the Llano Estacado of northwest Texas (staked plains) according to Hafsten (1961). Bryant (1969) suggested that based on pollen profiles, the present Chihuahuan desert area around Del Rio, Texas (430 m) was a pinyon woodland. Wells (1966), ADAMS —JUNIPERUS ASHEI 197 1977] HYPOTHETICAL PLEISTOCENE PLUVIAL VEGETATION 1I0-15,000 bp OZARK MTNS. BOREAL SPRUCE FOREST / ARBUCKLE MTNS. PINE-SPRUCE Ç | Š | ^4 : < | = o ! = / FOREST 9 | Lan? a! o0 =É | w0* AN? A ine RKL 14 Z P pA x E "4 | MIXED DECIDUOUS DAVIS MTNS. WOODLAND WITH b EDWARDS PLATEAU j CONIFERS x AA CHISOS MTNS. rad NA, [PINYON-JUNIPER __— , WOODLAND a of. * sierra M’ 3 DEL CARMAN A S. DEL BURRO A O IOO 200 300 A A A KILOMETERS A Z x GUATRO CIENEGAS Hypothetical Pleistocene pluvial vegetation, 10,000-15,000 B.P. based on Fic l pollen profiles and rat midden data from the literature. See text for discussion. using data obtained from rat middens from the Big Bend Texas region, con- cluded that the life zones descended about 800 m for pinyon-juniper (J. pinchotii in that case), allowing the advance of pinyon-juniper into most of the present desert region between Big Bend and Del Rio. Another important fact has been the recent discovery (D. H. Riskind, pers. comm.) of J. ashei, J. pinchotii, J. flaccida, and J. scopulorum from the Sierranas del Burro (Fig. 11). Since typi- cal J. pinchotii has been found (Adams, 1975b) just south of the Sierra del Car- man (growing with J. ashei), it appears that the Sierranas del Burro may have been an important refugium or island point in the pinyon-juniper woodland. A mixed deciduous woodland with conifers is postulated in central Texas (Bryant 1969; based on a pollen profile). 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE 4. SNK tests for 15 morphological characters with | F ratios greater than w: in ANOVA. SNK tests were run at tP = dpi dede = 1.58, Fou = 1.90 (df = 27/380). Any two populations not underlined by ommon line are significantly different. Popula- tions are listed in end order ft 3 due means from largest to smallest. Ranges refer to the maximum and minimum means over all 3 for that character. FEMALE CONE DIAMETER (FDI), F = 9.3, no obs. for populations 24, 27, range = (8.91-6.4 mm) 155 1463 23 47 11 21 16 2 10 19 8 22 28 20 1 18 9 13 12 17 26 25 SEEDS PER FEMALE CONE (SPF), F — 9.7, no obs. for populations 24, 27, range — ( 1.69-1.01) 12 26 13 25 4 17 11 28 2 19 3 10 14 16 21 1 18 8 15 23 20 5 7 6 9 22 BLOOM ON FEMALE CONES (BLM), F = 1.2, no significant differences SEED AREA (SEA), F = 9.8, no obs. for populations 24, 27, range (27.1-13.6 mm’) 20 16 21 145 15 4 23 6 22 8 7 1039 19 2 28 11 18 1 17 13 12 25 26 SEED WIDTH/LENGTH (SER), F = 1.3, no 5 differences WHIP GLAND AREA (WGA), F 9.2, range (0.93-0.31 mm?) 12 25 11 13 3 26 23 17 56 28 14 7 20 16 21 4 18 1 22 24 95 Daora WHIP LEAF GLAND LENGTH/WIDTH (WGR), F = 4.4, range (2.5-1.5) 13 25 17 26 5 12 114119827 14 3 27 22 16 18 20 6 9 10 23 15 28 24 21 WHIP LEAF MARGINS (WLM), F = 5.0, range (2.3-1.9) 24 28 25 22 23 26 16 13 21 27 5 17 96 19 18 20 1443 15107112811 WHIP LEAF GLANDS PROTRUSION (WGP), F = 2.6, range (3.00-2.87 ) 1367 1011 15 19 20 21 23 25 18 12 17 22 28 16 8 9 14 2 24 13 4 5 26 27 WHIP LEAF GLANDS RUPTURED (WRP), F = 2.7, range (1.08-1.00) 1312111446789 102315 15 16 17 18 19 20 21 22 23 24 25 26 27 28 WHIP LEAF BLADE LENCTH/SHEATH LENGTH (B/S), F = 2.9, range (0.77-0.54) 24 25 2 26 9 12 11 1 27 28 14 15 16 1065 13 19 22 2021 4 7 3 8 18 23 17 WHIP LEAF GLAND LENGTH/SHEATH LENGTH (G/S), F = 16.7, range (0.41-0.22) 25 26 12 13 17 11 24 3 28 19 22 21 23 164 5 18 2096 14 2 108 1 7 27 15 SCALE LEAF LENGTH (SLL), F = 4.1, range (1.74-1.43 mm) 28 27 24 25 197 18 16 1 22 15 26 14 21 204 5 23 11 139 17 1232 8 6 10 SCALE LEAF LENGTH/BRANCH WIDTH (L/B), F = 4.1, range (1.43-1.15) 927 17 1519 185 167 201 14 8 2223 4 21 25 28 11 3 2 13 6 10 24 12 26 BRANCHING ANGLE (BAN), F = 9.5, range (55.2-39.9 degrees) 12 26 25 1024 13 27 207 17 518112819 1694238 156 14 222112 3 1977] ADAMS JUNIPERUS ASHEI 199 Life zones were pushed southward and compressed during the Wisconsin pluvial ( Dillon, 1956), but how far they were extended into Mexico is not well known. Additional rat midden and pollen profiles are needed in northern Mexico and the Mexican plateau region. A study by Meyer (1973) in the Cuatro Ciene- gas basin (Fig. 11) revealed no changes in the pollen profiles during the past 30,000 years. He concluded that there was no evidence for pluvial nor hypsi- thermal ( Deevey & Flint, 1957) periods at Cuatro Cienegas during the time se- quence studied. This agrees with Dillon (1956: 174) who shows a considerable compression of life zones from Nebraska to south Texas but few differences past northern Mexico. It seems likely that any generalized Mexican refugium must have been in northern Mexico. One other point that seems relevant is that Wells (1966) mentioned that the pinyon found in the rat middens in the Big Bend area contained consistently 2-needled fasicles suggesting that the pine involved may not have been the predominately 3-needled Pinus cembroides Zucc, but perhaps Pinus cembroides var. remota E. L. Little. Pinus cembroides var. remota now persists on the Balcones escarpment of the Edwards Plateau (near population 14, Fig. 1) about 300 km to the cast of the fossil site. I have recently examined a herbarium specimen of J. ashei from eastern Brewster County, Texas and have indicated this location in Fig. 1 (dashed lined population, about 150 km west of population 26). This new population is just north of Wells's (1966) Maravil- las Canyon rat midden site. Perhaps his juniper twigs should be reexamined for the presence of J. ashei. In any case, this western-most disjunct population of J. ashei seems to be of the same relict nature (on preliminary morphological examination) as populations 12, 13, 25, and 26. Although I have previously considered populations 12, 13, 25, 26, and 17 to be ancestral ( Adams, 1975a), one might ask why these might not be advanced, with the central Texas-Ozark populations being ancestral. Examination of Table 4 reveals the significant morphological differences between populations 12, 13, 17, 25, 26, and the other populations. The following characters show significant differences: female cone diameter (smaller in 25, 26, etc.); seeds per cone (generally more in 25, 26, etc.); seed area (smaller in 25, 26, etc.); whip leaf gland area (larger in 25, 26, etc.); whip leaf gland length/width (more elongated in 25, 26, etc.); whip leaf gland length/sheath length (larger in 25, 26, etc.): and branching angle (larger in 25, 26, etc.). Reviewing the Sabina section of Juniperus in North America, it seems that some of these character states are rather unusual and are likely advanced (rather than primitive). Advanced charac- ter states (central Texas-Ozarks) are: larger female cones concurrent with fewer seeds (just the opposite found in most of the junipers); whip leaf gland area small (whip leaf gland area is generally large in junipers where the glands are visible); whip leaf gland length/width close to 1 or 1.5 (J. ashei is unique in the genus, so far as is known, in having raised, round glands), the more elon- gated glands (populations 25, 26, etc.,) are definitely the more primitive type; and whip leaf gland length/sheath length (almost always large in Juniperus, except the central Texas-Ozark J. ashei). Advanced and primitive states are not known for two characters: seed arca and branching angle. Overall, the characters expressed in central Texas and the Ozarks are generally unusual in occurrence 900 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 2 MISSOURI e 3 e 3 (3) 1 ARKANSAS 2 XARKANA TEXAS 100 200 100 90 | = KILOMETERS | 8 CHEM. ‘SR, ASHEI-SALTILLENSIS MORPH. SR, ASHEI-SALTILLENSIS URES 12-13.—12. The contoured F-1 weighted morphological similarity of each popu- lation of J. ashei to J. saltillensis, collected near Saltillo, Mexico. Juniperus saltillensis is thought to be closely related to the ancestral stock of J. ashei. Notice n P pp die tion from west Texas to the Mexico population (25). Contours: 1 — T. — —13. The contoured F-1 weighted terpenoid similarity of each population of 7 D to x “alte n- sis. The clinal trend seen with the morphology (Fig. is steeper in the terpenoids, and population 17 is 1 more closely related to the ancestral stock of J. saltillensis-]. ashei Contours: 1 — 0.17; 7 = 0.41. in Juniperus compared to the character states found in the southwest Texas- Mexico populations. Further evidence regarding the ancestral nature of popu- lations can be obtained by comparison of each population of J. ashei with its presumed nearest ancestor (Zanoni & Adams, 1976), J. saltillensis Hall. Although J. ashei probably did not descent from J. saltillensis, that taxon appears to bear the closest morphological and terpenoid similarities to J. ashei of any in North America. In Fig. 11 I have constructed differential similarities of each popula- tion of J. ashei to a sample of 15 trees of J. saltillensis from near Saltillo, Mexico. ANOVA was performed on 29 data sets (28 J. ashei populations and 1 J. saltillen- sis population) to determine a set of F-1 weights. Similarity measures were cal- culated as outlined before, then each population of J. ashei was contour mapped showing the change (differential) in similarity to J. saltillensis (the geographical source of this taxon is not important for obtaining the similarities and is not shown on the maps). This method of "differential similarity" should prove very useful in the analysis of the interaction of two species across a geographical area. Figure 12 is based on 15 morphological characters (female cone color omitted, F — 0.88), F-1 weighted. Notice that the highest similarity to J. saltillensis is from the Mexico population (25), followed by populations 26, 12, 13, and 17. The knife edge break previously seen (Fig. 4) between populations 12, 13 and 11, 14 is quite widened in this analysis with a cline from populations 12 to 10. The Post population (24) bears some similarity, but part of this similarity may be due to environmental factors. The terpenoids of J. ashei are interesting evolutionarily because there is a 1977] ADAMS JUNIPERUS ASHEI 20] greater shift toward the predominance of a single compound (camphor, see von Rudloff, 1968; Adams & Turner, 1970) than in any other member of the genus. In populations of central Texas camphor averages about 75% of the total oil (2 hr. extraction) whereas the divergent populations average about 60%. Juniperus ashei has by far the simplest oil mixture of the North American junipers, and this seems to be an advanced character state of specialization. The central Texas populations are particularly low in the sesquiterpene oxygenated compounds such as elemol, clemol-acetate, and a, B and y-endesmols. Larger quantities of these compounds are the rule in the rest of the junipers and conifers in general (see von Rudloff, 1975). Differential similarities, based on ANOVA (28 J. dei populations plus 1 J. saltillensis population) and using 68 terpenoids F-1 weighted, reveal (Fig. 13) a pattern almost identical to the differential similarities for the morphologi- al characters (Fig. 12). These similarities indicate that the divergent populations (25, 26, 12, 13, 17) bear a stronger affinity to J. saltillensis than the central Texas-Ozark populations (lest the reader be suspicious of mixed sampling in population 25, etc. I should note that these divergent populations clustered strongly with the central Texas type when an OTU of J. saltillensis was added to the matrix set, and intrapopulational cluster analysis of each of the 28 popu- lations of J. ashei revealed no other taxa as would be the case in mixed species samples). Thus we see that in considering a fairly large set of characters (15 morphological and 68 terpenoids), the dominant theme is for the divergent popu- lations to be progressively more similar to J. saltillensis. It should be noted that J. saltillensis is not conspecific with J. ashei (Zanoni & Adams, 1975, 1976). In fact, several characters found in J. saltillensis (curved terminal whips and beady scale leaves) have not been found, even in the relict populations, in J. ashei. Al- though relict hybridization could not be conclusively ruled out at present, it seems unlikely since we have no direct evidence that the two taxa have been sympatric, and several distinguishing characters of J. saltillensis have not been found in divergent J. ashei plants. It would appear that the most probable hy- pothesis at present is that J. ashei and J. saltillensis had a common ancestor ( Ter- tiary?) in the Sierra Madre Oriental. Juniperus ashei differentiated and migrated northeastward to the exposed limestone outcrops (Edwards Plateau, Arbuckles, Ozarks, etc.), while J. saltillensis adapted to the drier, interior portion of the Sierra Madre Oriental. During the Pleistocene ice advances, J. ashei may have become extinct in Missouri, Arkansas, Oklahoma, and most of central Texas as depicted in Fig. 14. During the same period, J. ashei probably expanded westward into the current Chihuahuan desert ( Wells, 1966; Bryant, 1969), but not as far south as Cuatro Cienegas ( Meyer, 1973). Migration west of the Sierra del Carman was also pos- sible since the species is currently found at the top of a pass (La Cuesta) just south of the Sierra del Carman. Whether J. ashei could have crossed the high plateau around Alpine and Marfa (1,500 m) is not known, but suitable habitat — was probably available for colonization in the Presidio area. With this model, populations of J. ashei would be forced to extinction in central Texas, Oklahoma, Arkansas, and Missouri. The subsequent recolonization could then take place ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 WISCONSIN DISTRIBUTION OF J. ASHE! 10-15,000 bp OZARK MTNS > a / ARBUCKLE MTNS. g ! n 8 | MA < ' Ü o | u LOCAL EXTINCTIONS 9 | OF J ASHE! R 7 — = E. A — T Z aS A i 2 | DAVIS MTNS à | BALCONES Ø EDWARDS PLATEAU "| ESCARPMENT Z x JSURVIVAL (RELICT, CHI Pd ? — — RETREAT TO PINYON MP EDULIS REMOTA: DEL CARMAN JUNIPER (J. ASHE|, PINCHOT II) ? A WOODLAND 7 o 100 200 300 Gmm u "gd KILOMETERS x CUATRO CIENEGAS NSS = REMNANT, ADAPTED TO MORE MESIC ENVIRONMENT Y = ANCESTRAL TYPE POPULATIONS, LOW CAMPHOR TYPE FicurE 14. Possible Wisconsin distribution of J. ashei, 10,000-15,000 B.P. Following ce of subalpine and montane speci Fig. 11), J. ashei populations may have gone the adv extinct north of the Edwards Plateau. See text for discussion. — according to Fig. 15 over a very short period of time (hundreds of years?) from some population in central Texas that may have gone through a selection “bot- tleneck,” perhaps coupled with genetic drift. This “relict” population would have had considerably more camphor in the oil (as a plant defense?), more roundish glands, larger female cones, fewer seeds (therefore a higher pulp to seed ratio for bird dispersal), and a more lax foliage (smaller branching angle) which seems to be associated with more mesic species. The rapid recolonization of limestone outcrops (Fig. 15) could then lead to a uniform taxon from central Texas through ADAMS —JUNIPERUS ASHEI 203 1977] POST GLACIAL MIGRATION AND DISTRIBUTION OF J. ASHEI OZARK MTNS. / / ARBUCKLE MTNS. o ! ^ < | A I: T AM ? WwW ' — = ' < i 3 J A z M =~ ' 5 "s SSS | DAVIS MTNS. j : J h EDWARDS D É PLATEAU 100 200 300 KILOMETERS x CUATRO CIENEGAS = REMNANT, ADAPTED TO MORE MESIC ENVIRONMENT DURING THE PLUVAL PERIOD FA = PRESENT DISTRIBUTION OF RELICT POPULATIONS Possible post-glacial migration to attain the present distribution of J. ei. a more mesic environment may ickly ribution (see Fig RE 15. The remnant (high camphor type) adapted to expanded during the hypsithermal to reach the present dist line shows the pluvial distribution of the ancestral type (lower camphor ) Fig. 14). the Ozarks. Although this would explain the observed patterns, many uncer- tainties remain. For instance, Dillon (1957) argues that the boreal forest ele- ments were merely mixed with the present floral components in the southern states. Graham (1973) feels that most central-southern communities incorpor- ated boreal elements (e.g., spruce) but retained the general character of the original vegetation. If small pockets of J. ashei did persist during the full glacial, 904 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 4 5 _ ô 6 MISSOUR 3 A 5 7 280 3 3 7 KLAHOMA RKAN SA 6 e4 ° POST 927 2 (O TEXARKANA 240 5 : ° 6 4 3S "s ^ : > q 6 $12 5.219 eg TEXAS 26, 1B Ag 7 eS ° 5 10Q_ 200 a 1 ROTER 5 X KILOMETE 34 5 —5432 2 3 4 ey 5 AVG. SIMILARITY, TERPENOIDS CPV, TERPENOIDS 5 6 3 2 18) [MISSOURI N — (9 MISSOURI OKLAHOMA Ñ \ f) ° 28 N 2 6 Ne 2 1 3 S 6 ARKANSAS) TEXAS 5 TEXAS 3 04 4 di 4 e27 3 e27 POST TEXARKANA POSTA N n 4 3 1 5 Q 2 S 3 p 21 6 N 266 IE. e9 1 5 Y 7 2 XJ Q 190 2945 2 16 2 O 100 200 25e TAE N KILOMETERS AVG. SIMILARITY. MORPHOLOGY CPV. MORPHOLOGY 3 4 4 3 2 1 7 O MISSOURI 5N en) MISSOURI AHQMA KLANONA 2 ge|3 *| Ú 289 |3 240 T S RKAÑ S TEXAS ANSAS 7 6 1 6 2 2 e4 5 e4 3 e27 s e27 TEXARKANA| > POST: XARKANA| Á e 21 V? e s< ^ 3 4 45 e7 4 12 1 e 1e8 4 26e 9 e9 3 O 100 200 14 %7 © 100 oo 2 — KILOMETERSN, Doe MEX KILOMETERS 2 3 4 5 3 2 AVG. SIMILARITY, PEROXIDASES |. CPV, PEROXIDASES 1977 ADAMS— JUNIPERUS ASHEI 20 cC one might find some evidence of this based on intrapopulation variability, with the smaller Pleistocene relictual populations having less variability than larger ( south-central Texas) populations. INTRAPOPULATIONAL VARIABILITY Mean similarity (Sr) within each population (15 trees) for 152 terpenoid characters (W = 1) shows high average similarities in the Ozarks and central Texas (10, 11, 15) and low similarities at New Braunfels (17), Post (24), and Mexico (25). The divergent populations (12, 13, 17, 25, 26) tend to be a little less uniform, although Post (24) is also quite variable. Populations that showed the major trend of the terpenoids (Fig. 3) tended to be uniform. Examination of the homogeneity of the similarities was accomplished by computing the stan- dard deviation of the mean similarity and dividing by the mean similarity of that population for normalization ( CPV). 'The most homogeneous similarities are in the Ozarks (2, 3) and central Texas (10, 15, 7), whereas the least homogeneous are New Braunfels (17), Brady (20), and Post (24). The populations which showed the highest similarities are the most homogeneous except for population 20 (Brady). The low similarities and lack of homogeneity at New Braunfels seems to be due to the interaction between relict and modern genotypes. One might question if the population at Post (24) is hybridizing with sympatric J. pinchotii trees but notice the close ordination of 24 with the central Texas J. ashei (Fig. 3). Examination of the intrapopulation phenogram revealed no major groups within any population. Analysis of 16 morphological characters (W = 1) shows the highest similari- ties in central Texas (7, 10, 18) and the Ozarks (1, 3), with lowest similarities in the relict populations (12, 13, 26, 17, 25) and at Texarkana (27). Two small island populations (27, 28) both show considerably lower similarities in their morphology than they did in their terpenoids, whereas Post (24) is more medial in its morphological similarities than with the terpenoids (Fig. 16). The CPV Ficunks 16-2 16. Average similarity (Sr) within each population. (15 trees) based on 152 equally ide: terpenoids. Most populations had high internal similarities with the n pu of Be ancestral populations (12, 13, 17, 25, 26) and the Post population (24). Contours: 1 = 0.78; 7 = 0.87 7. Contoured coefficient of phenetic variation (CPV) of the und 7 1 0 E general the populations with high intrapopulation similarities were homogeneous (low CPVs) and vice versa, except for 55 20 which ler: Ie similarities and was not so Duros neous (high CPV). Contour Í 0.30; 7 8 verage aagi within each population based on 16 dai de d bc em sharik ters. Note that the ancestral populations are of generally lower internal similarities com- pared to high la ie throughout central Texas. The small MD oi at Texarkana and northeastern Oklahoma are morphologically quite variable. Contours: 1 = 0.85; 7 = 0.91 —19, Contoured coefficient of phenetic variation (CFW) m die average Berg ages inn ate The CPV seems highly negatively correlated with the mean Sr except for popu- lations 19 and 20 which are not very homogeneous. Contours: 1 — 0.28; = 0.60.—20. Contoured average similarities of 23 equally weighted 8 The Junction pation (10) had the lowest similarities along with Post (24) and the Arbuckles (4). Contours 9 ~ ] = 0.54; 7 = 0.96.—21. Contoured coefficient of phenetic Patton of the peroxidase simi- larities gives an almost identical pattern as seen in the average similarities (Fig. 20). Con- tours: 1 = 0,13; 7 = 0.32. 906 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 gives a fairly similar pattern of homogeneity except for populations 19 and 20, which, although very typical (Fig. 4) and of high average similarities, are not very homogeneous. This same phenomenon was seen with the terpenoids (Figs. 3, 16-17) for populations 19 and 20. The Post (24) population is somewhat more homogeneous in its morphology than its terpenoid's similarity. With the excep- tion of populations 24 and 27, one notices that for each of these four statistics, the populations generally present some trend of variability which is correlated with either regional differentiation or proximity of one population to another (the case for 19 and 20). All 23 peroxidase electromorphs were subjected to the computation of aver- age similarities within population and CPVs, as with the morphology and ter- penoids. Only the 15 populations marked with an asterisk in Figs. 20 and 21 have isoperoxidases analyzed. The average similarities within populations for these 23 isozymes show the Ozark population (Fig. 20) to be quite similar (0.97-0.80), while the Junction, Texas population (10) has the lowest average similarity (0.47). A surprising aspect of these average similarities is the low average similarity found in population 10 (Junction, Texas). It is interesting that 3 different peripheral populations (1, 27, 24) show the whole range of vari- ation from little to large amounts to intermediate variability. The CPV (Fig. 21) of these 23 peroxidases reveals that those populations that are highly similar are generally most homogeneous and vice versa. A combined total of 43 iso- enzymes has been analyzed by Kelley & Adams (1977b), and the results are comparable to those shown in Figs. 20 and 21. However, the addition of 14 al- cohol dehydrogenases and 4 esterases to the analysis seemed to have produced a slightly less mosaic pattern in central Texas. The pattern obtained from the isoenzymes is quite different from either the morphology or terpenoids for in both of those analyses, population 10 appeared to be quite uniform and homogeneous and the relict populations consistently displayed high to medium variability. It seems apparent that whatever vari- ability the peroxidases are indicating, it is not directly related to variability in the morphology nor terpenoids. Of course, it is possible that the variation seen in peroxidases is below the level of selection and merely represents "random noise." Until more information is gathered on the selection value of various electromorphs, we can only speculate. The patterns of variability seem to give us a few clues as to whether the dis- junct populations are of recent origin or relicts of the advanced high camphor types. However, presently it is difficult to make generalizations about popula- tional variability versus founder's effect, “bottlenecks,” relictness, etc. since different character sets give somewhat (to vastly) different answers, and one could get the same observed pattern depending on time, microselection inten- sity, or site variability. CONCLUSION Treating peroxidase data as qualitative taxonomic chemical characters did not appear to be feasible. This is likely due to the lack of homology, intense microhabitat selection or random (neutral) variations in the electromorphs. 1977] ADAMS—JUNIPERUS ASHEI 207 The use of these peroxidase electromorphs for the analysis of intrapopulation variability could not be readily evaluated due to the mosaic pattern produced. It appears that chemosystematists will need more detailed biochemical informa- tion about the nature of isoenzymes, and their genetic control in taxa to be studied. The most probable center of origin for the modern (high camphor) popula- tions of J. ashei seems to be in central Texas, perhaps near Brady (20) or Bur- net (19). These populations showed considerable variability (high CPVs). yet these populations are quite similar to the rest of the modern J. ashei popu- lations. Northward migrations of birds during the spring carrying juniper seeds could have (re)colonized limestone outcrops in Arkansas, Oklahoma, and Mis- souri in a span of a few hundred years. This could lead to the highly uniform pattern observed in the morphology and terpenoids from central Texas to the Ozarks. Predominately southerly winds during pollination may have been im- portant in maintaining the north-south split in west Texas as well as the relict population at New Braunfels, Texas (17). However, it is possible that the popu- lations persisted throughout the pluvial periods and failed to diverge due to either a lack of variability, the relatively short time span involved, or intense se- lection for the modern phenotype. Whether the modern populations of this taxon invaded the limestone outcrops in the Tertiary or during the Pleistocene will probably not be known until some pollen or macrofossil (rat midden) data has been analyze in the disjunct popu- lations of Arkansas, Missouri, and Oklahoma. The differential similarity of J. ashei population to J. saltillensis from Saltillo, Mexico shows a clear trend of past (Pleistocene) migration from northern. Mexico. The northern Mexico Sierra Madre Oriental seems a likely site for the origin of both J. ashei and J. saltil- lensis, perhaps from a common ancestor. This study presents additional evidence that selection may be more impor- tant than gene flow (Ehrlich & Raven, 1969) in the maintenance of species. In J. ashei we have found that populations with disjunctions of 200-300 km, and a trivial chance for gene exchange, were very similar to other populations cover- ing 1,000 km of range (cf. Ozarks and central Texas populations). Yet popula- tions which are in close (almost continuous) proximity have maintained either ancestral or modern patterns in spite of potentially large amounts of gene flow. (New Braunfels and populations to the north and west, and the relict/modern populations of west Texas) LITERATURE CITED ApAMs, R. P. 1970. Seasonal variation of ore e; constituents in natural populations of Juniperus. Men Sudw. Phytochemistry 9: 397-402. 1972. Numerical analysis of some common errors in chemosvstematics. Brittonia 24: 921. 19754. Gene flow versus selection pressure and ancestral differentiation in the n of species; analysis of RED variation of Juniperus ashei Buch. using ter- qas data. J. Molec. Evol. 5: 177-1 1975 5b. Numerical- e is studies of infraspecific variation in Juniperus — Sudw. Biochem. Syst. Ecol. 3: 71-74. 1975c. Statistical character weighting and similarity stability. Brittonia 27: 305- - 316 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 B. L. Turner. 1970. 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Vol. Academic Press, New York. KELLEY, W. A. 1976. Isoenzyme studies in 7 sn Ph. D. thesis, Colorado State Univ., Ft. `Collins, Colorado. & R. P. Ap AMs. 1977a. Preparation of extracts from juniper leaves for electrophore- sis. Phytochemistry 16: 513-516. & —. 1977b. Variation in isozymes from natural populations of Juniperus. Rho- dora (in — KiNG, J. E. 1973. Late Pleistocene palynology and biogeography of the western Missouri Ozarks. Ecol. Monogr. 43: 539-565. McCrunrE, J. W. & R. E. ALSTON. 1966. A chemotaxonomic study of Lemnaceae. Amer. J. Bot. 53: 84$ ot )-859 MEHRINGER, P. J., Ja. J. E. Kinc & E. H. Linpsay. 1970. A record of Misi a Ses ve RE tation s fauna from the Ozarks of western Missouri. Pp. 173-184, in W. Ir. & x jones, Jr. (editors), Pleistocene and n Environments of the Psi T Great Plains. "Univ. of Kansas Press, Lawrence, Kans 1977] ADAMS—JUNIPERUS ASHEI 909 Meyer, E. R. 1973. Late quaternary paleoecology of the Cuatro Cienegas basin, Coahuila, Mexico. Ecology 54: 982-995. Mooney, H. A. & W. A. EMBODEN. 1968. The relationship of terpene composition, morphol- ogy, mee distribution of populations of Bursera microphylla (Burseraceae). Brittonia 20: 44- SMITH, D. vh & D. A. Levin. 1963. A chromatographic study of reticulate evolution in the 3 Asplenium complex. Amer. J. Bot. 50: 952-958. SmitH, R. H., R. L. PELoQuiN & P. C. Passor. 1969. Local and regional variation in the monoterpenes of ponderosa pine wood oleoresin. U.S.D.A. Forest Serv. Res. Pap. PSW- 56: 1-10. VAN Devener, T. & J. E. KING. 1971. Late Pleistocene records in western Arizona. J. Arizona Acad. Sci. 6: 240-244. VON ee E. 1968. Gas-liquid chromatography of W Part XVI. The volatile yil of the leaves of Juniperus ashei Buchholz. Canad. J. Chem. 46: 679-683. 197 5. Volatile leaf oil a ed sis in chemosystematic mo of North American coni- fers. Biochem. Syst. Ecol. 2: 58. WkLLs, P. V. 1965. Scarp ded: transported grasslands, soils, and concept of grass- land : P ps eee Region. Science 148: 246-249. pleistocene vegetation and degree of pluvial climatic change in the bs ted p. y Science 153: 970-975. 1970. Postglacial vegetational history of the Great Plains. Science 167: 1574- 1582. have Desert. Science 155: 1640- WHITEHEAD, O. R. 1972. Approaches to disjunct populations: the contribution of paly- nology. Ann. Missouri Bot. Gard. 5 25-137. WILLIAMS, W. T., B. Dare & G. | 3 1971. Two outstanding ordination prob- lems. Austral ]. Bot. 19: 251-2 W RIGHT, H. E., Jr. 1970. Yekretatioñal history of me Ce veo E 5 Pp. 157-172, in W. BEncER. 1967. Late 1 iai history of coniferous woodland in the Mo- 16 Dort, Jr. & J. Knox Jones, Jr. (editors), Pleist tocene and Recent Environments of the Central ew Plains. d . of Kansas Press, Lawrence, Kan ZANONI, T. A. P. ApAN 1975. The genus Juniperus ( Cupressaceae) in Mexico and ea REI and morphological analysis. Bol. . Bot. México 35: 6 1976. The genus Juniperus nM in - and Bu temali: — aiid chemosystematic analysis. Biochem. Syst. Ecol ZAVARIN, E. & K. SNAJBERK. 1973. Variability of wood monoterpenoids p Pinus aristata. id Syst. Ecol. 1: 39-44. THE ORDER CENTROSPERMAE' ToM J. MABRY2 ABSTRACT Perhaps no other order of flowering plants of its size is as well investigated morphologi- cally, ultrastructurally, and chemically as is the Centrospermae. The betalain pigment dis- based on 11 core families, including all 9 betalain families: Aizoaceae, Amaranthaceae, Basel- hytolac laceae, Cactaceae, Chenopodiaceae, Didiereaceae, Nyctaginaceae, Phytolaccaceae, and Por- tulacaceae, as well as two anthocyanin families: Caryophyllaceae and Molluginaceae. goi smaller betalain taxa (including Gisekia. Halophytum, Hectorella, and Dysphania) whic sometimes treated as independent families or as members of one of these 11 core families k trospermous families may have originated from a centrospermous na which lost the ability to produce anthocyanins and then subsequently gained the two or three steps required to produce betalains. Pollen morphology of centrospermous taxa and the widespread occurrence of C, photosynthesis in the Centrospermae are also discussed. Since all the review papers from a symposium on the “Evolution of Centro- spermous Families,” presented in July, 1975 during the XIIth International Bo- tanical Congress, Leningrad, USSR, have now been published (Mabry & Behnke, 1976a), a summary of our current views of the Centrospermae (or Caryophyllales) will suffice in this review. This account will emphasize the way our interpre- tations of the order have been shaped by molecular data. Since 1876 when Eichler (see Table 1) introduced the name for the order, the Centrospermae have always contained a core of about 8-12 families. Eichler (1876 and, in part, 1878) recognized most of what we now consider to be centro- spermous families including, for example, the Cactaceae (in the 1876 treatment). In the 100 years following Eichler's work, most systematists also included those families now generally recognized on the basis of molecular data as belonging to the order but often included additional ones (compare Tables 2, 3 and 4). The molecular data which bear upon our current treatment of the order are sum- marized in the following sections. SIEVE-ELEMENT PLASTIDS Of all the modern approaches for investigating the Centrospermae, none has, in my opinion, contributed more to our understanding of the circumscription of the order than the ultrastructural investigations of the sieve-element plastids. ' Portions of the research described here were supported by NSF Grant DEB 76-09320, NIH Grant HD-04488, and Robert A. Welch Foundation Grant F-130. Helpful discussions with, and in some 1 unpublished data from Profs. Martin Ettlinger, H.-D. Behn Walter Brown, H. Musso, F. Ehrendorfer, and Jerry Brand are gratefully 5 ? Department of Botany, The University of Texas at Austin, Austin, Texas 7871 ANN. Missouni Bor. Garp. 64: 210-220. 1977. 1977] MABRY—CENTROSPERMAE 211 TABLE I. Centrospermae (A. W. Eichler, 1876). I. Order: Oleraceae III. Order: Opuntinae l. Polygonaceae 6. Phytolaccaceae* 2 yctaginaceae* 7. Portulacaceae 3. Chenopodiaceae* 8. Aizoaceae* 4. Amaranthaceae* 9. cta , IL. Order: Caryophyllinae (? In this Order perhaps 5. Caryophyllaceae Begoniaceae ) * Betalain families. The discovery by Behnke that the nine core betalain families (see Table 4) well as the two core anthocyanin families (the Caryophyllaceae and Mollugin- aceae) contain ringlike inclusions composed of proteinaceous filaments of type (Fig. 1) not found elsewhere in the angiosperms (for current reviews, see Benhke, 1976a and in press) established that these eleven families represent the core centrospermous families. Behnke (in press) defined the sieve-element plastids which are unique to the Centrospermae as belonging to the P-III sub- type. It is my current view that the presence of the P-III subtype sieve-element plastids in a taxon which classical data suggest might be centrospermous (see Eckardt, 1976 for comments on centrospermous characters) establishes that it belongs to the Centrospermae. C, PHOTOSYNTHESIS IN THE CENTROSPERMAE It is interesting that among the dicotyledons which have the C, photosynthetic pathway (the Kranz syndrome), 7 of the 11 families (Table 5) and about 85% of the genera have been reported among the Centrospermae (Walter Brown, private communication). It is perhaps significant that the Phytolaccaceae, which most workers consider to be the basal family of the order, exhibits only the C; TABLE 2. Caryophyllidae ( Cronquist, 1968). I. Order Caryophyllales II. Order Batales 1. Phytolaccaceae* (incl. a 1. Bataceae Agdestidaceae*, Barbeuiaceae ;yTO- III. Order 1 stemonaceae, Petiveriaceae", Stegno- l. Polygo spermaceae* IV. Order B us N ginaceae* 1. Plumbaginaceae E d. c5 po e > e O lo ^ — — — = = — — > ea Caryophyllaceae (incl. Illecebraceae) Portulaca eae Basellaceae * pene. (incl. Dysphaniaceae*, Halophyt 11. Amaranthaceae | - SO = > * Betalain families. 912 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE 3. Caryophyllales, Polygonales and Plumbaginales (Takhtajan, 1973). 27. Ordnung. Caryophyllales 10. Basellaceae* 1. Phytolaccaceae* (incl. Achatocar- 11. Didiereaceae* paceae, (ERE aliu, Bar- 12. Halophytaceae* beuiaceae, Petiveriaceae*, Steg- 13. Hectorellaceae* nospermaceae,* excl. (?) Rhab- 14. . (incl. Illecebra- odendro 2. 1 15. Vivianiac ceae 3. 16. Amaranthaceae* 4. sana d 17. Che sci" acl (incl. Dysphania- 5. Molluginacene. (incl. Gisekiaceae* ) 6. Aizoaceae* 98. Ordnung. Polygonales 7. Tetragoniaceae* Polygo 8. Cactaceae* 29. Gace “i us 9. Portulacaceae* Plumbaginaceae (incl. Armeriaceae ) * Betalain families. pathway. Although the C, pathway probably represents a derived condition as taxa of the order radiated into xeric and other high light intensity habitats, it is likely that some form of preadaptation for the Kranz syndrome exists through- out the order. This preadaptation has permitted repeated and independent evo- lution of the syndrome at various times and in numerous genera in at least seven families of the order (Walter Brown, private communication). The only other dicotyledonous families exhibiting the C, pathway are the Boraginaceae ( Helio- tropium in part), Compositae (7 genera), Euphorbiaceae (Chamaesyce), and Zygophyllaceae (3 genera It should also be noted that the CAM (crassulacean acid metabolism) photo- synthetic pathway occurs in members of the Cactaceae and Aizoaceae; this path- way, although anatomically, functionally and phylogenetically distinct, does uti- lize enzymes of the Kranz syndrome. Although CAM plants are photosynthetically TABLE 4. Order Centrospermae* or Caryophyllales (modified here from Mabry, 1976). (Taxa d P-III subtype sieve-element plastids. ) SUBORDER CHENOPODIINEAE" (BETALAIN Nyc FAMILIES ) Phytolaecacese (incl. Achatocarpaceae, Ag- ae Aizoaceae (incl. Tetragoniaceae and pos- eae, Petiveriaceae, Stegnosperm- sibly Gisekiaceae k ; 5 (incl. Hectorella) Basellaceae SUBORDER CARYOPHYLLINEAE (ANTHOCYANIN š FAMILIES ) .actaceae D dde (incl. Dysphaniaceae ) ereaceae Haloph ytaceae Caryophyllaceae Molluginaceae (excl. Gisekia) a Certain families which on occasion. have been treated as members of the Centrospermae but are now known to contain neither the sieve-element plastids (subtype P-III) (see Behnke, 1976a, 1976b, in press) nor betalains are excluded from the order: Polygonae eae, 8 Fouquieraceae (Behnke, 1976b), Frankeniaceae (Behnke, 1976b), Rhabdodendron (Behnke E ge: = mains to be Mane for pigments), Vivianiace: m ne hnke & Mabry, Sur Theligonaceae ( 1975), and Bataceae and Gyrostemonaceae. So either anthocyanin r betalains have E El in the latter two "families both of which contain . (see Goldblatt et al., 1976 for recent com- ments gnt the status of these two families). Whether or not such betalain taxa as Petivera and Agdestis (Behnke et al, 1974), Halophytum (Hunziker et al., 1974), Gisekia (Mabry, 1 28 c 1976), Dysphania ( Mabry & Behnke. 1976b) and 5 (Mab ry, preliminary results) should h be treated as families in the suborder Chenopodiineae or as members of one of the core betalain families is not potu ie 1977] MABRY—CENTROSPERMAE 913 TaBLE 5. Kranz and CAM Photosynthesis* in the Order Centrospermae" (data from Walter wata] private communication, 1976). Suborder Chenopodiineae Suborder Caryophyllineae (Betalain Families ) (Anthocyanin Families ) Aizoaceae Kranz and CAM Caryophyllaceae Kranz Sesuvieae Kranz ychnideae 3 Gisekiaceae Kranz Polycarpeae Kranz (1 genus) Petra 1 C. Paronychieae Cs Amaranthac Kranz (11 genera) Diantheae Cs Basellaceae C; (3 genera Alsinea C; Cactaceae CAM Sperguleae y Chenopodiaceae Kranz (31 of 67 gen- Molluginaceae Kranz (2 genera) Dysphaniaceae 3 Nyctaginaceae Kranz (3 genera) Phytolaccaceae Ja hatocarpaceae Ca Agdestidacea ' Petiveriaceae C: Stegnospermaceae C: Portulacaceae Kranz (1 genus) a bagel = C, photosynthesis; CAM = crassulacean acid met holism pathways examined: Didiereaceae, Hectorellaceae and Halophytaceae. (Added in proof: Didiereaceae and W are C,.) inefficient, they are efficient at conserving water (see, for example, Winter, 1974) since they, unlike C; and C, plants, have their stomata open only at night. In contrast, Kranz plants have probably been selected for efficient photo- synthesis since they, unlike C, plants, have evolved an anatomy and enzymatic system which permits them to provide the Calvin cycle with high levels of CO. in an oxygen-deficient atmosphere. In the presence of low oxygen concentrations the enzyme which fixes CO, in the Calvin cycle, ribulose diphosphate carboxylase oxygenase, is free to function strictly as a carboxylase; therefore under these con- ditions and with high CO. levels and high light intensities, CO» fixation is ap- parently maximized. Thus the reasons for selection of the Kranz syndrome in the Centrospermae (and elsewhere) may be associated in part with evolution in habitats of high light intensity, which, of course, includes many of those which are xeric. Other factors such as salinity also may be important for the selection of C, and CAM photosynthesis. In any case, the widespread occurrence of the Kranz syndrome in both betalain and anthocyanin centrospermous families and its sporadic and limited occurrence in other dicotyledons supports current treat- ments of the Centrospermae (see Table 4) and provides additional evidence that the Centrospermae is an old, independent evolutionary line in the angio- sperms. PicuENT DicHorouay AND DNA-RNA HYBRIDIZATION DATA ron CENTROSPERMOUS FAMILIES Higher plants usually contain vacuolar red and yellow pigments which are either anthocyanins or betalains (Fig. 2); however, so far as known, the two types of pigments never occur together in the same plant or even in species 914 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 (a) (b) (c) globular crystalloid polygonal crystalloid no crystalloid Caryophyllaceae Chenopodiaceae Cactaceae Amaranthaceae Didiereaceae | Tetragoniaceae Aizoacea Molluginaceae Limeum Hectorellaceae Portulacaceae Nyctaginaceae Stegnosperma "d Phytolaccaceae Ficun The P-III subtype sieve element plastids which are characteristic for the Cen- trospermae always contain a ring-shaped bundle of protein filaments with either globular (a) or polygonal b) oni crystalloids or no crystalloid at all (c). I gratefully thank Prof. H Behnke for this figure which illustrates the distribution of these three modifications of P- TH subtype in the Centrospermae. of the same family. Of these two types, anthocyanins are much more wide- spread, indeed, they account for most flower pigments in higher plants (for recent reviews of anthocyanins, see Timberlake & Bridle, 1975; Harborne, 1967). In the 1960s it became clear that most centrospermous families contained an entirely new class of red and yellow pigments, designated in 1966 as the “betalains” (Mabry & Dreiding, 1968; for more current reviews, see Mabry, 1973, 1976; Mabry, Kimler & Chang, 1972; Piattelli, 1976). Although we know today that betalains also account for some of the orange and red pigments in many mush- rooms, notably species of Amanita ( Dópp & Musso, 1973, 1974; von Ardenne et al., 1 , these nitrogenous pigments have not yet been reported outside the Centrospermae among angiosperms. It is this restricted distribution of betalains as well as their being mutually exclusive with anthocyanins that makes them interesting as phylogenetic markers. 1977] MABRY—CENTROSPERMAE 215 Red Yellow + + HO N coo? ^N coo? | HOOC CNI “COOH Hooc co H H À max 537 Ara ^74 OH HO C HO C Mar Oe Zz Zz OR OH OH OR A max 534 Asas 477 SURE The visible absorption maxima of typical red and yellow betalains (top row), which are found only in phyletically related cen * families rea mushrooms, are similar to those for some red and sr ds anthocyanins (bottom row). Anthocyanins account for pigments in most plants, including members of two centrospermous M uA the Caryophyl- laceae and Molluginaceae. BIOGENESIS OF BETALAINS It is the biogenesis of betalamic acid and its subsequent condensation with amino acids and amines which appears to be significant among angiosperms for centrospermous plants. It now appears that the 4,5-extra diol cleavage of L-dopa (Fig. 3) can lead to betalamic acid (Fischer & Dreiding, 1972; Impellizzeri & Piattelli, 1972; Chang et al, 1974) in the Centrospermae and the mushrooms (Musso, private communication) and to stizolobic acid in the anthocyanin-con- taining Leguminosae (Ellis, 1976) and also mushrooms (Saito et al., 1975, 1976; Musso, private communication). Yet the conversion of the cleaved product to betalamic acid and its conversion into other betalains is known for the Centro- spermae and mushrooms only. Whether or not these different groups of organ- isms utilize the same enzymes to synthesize betalains is not known. PHYLOGENETIC SIGNIFICANCE OF BETALAINS Among, angiosperms, betalains are known only for centrospermous families, and we use this unique character to circumscribe the suborder Chenopodiineae, order Centrospermae (see Table 4). Whether or not all systematists accept this particular subordinal treatment is not important; it is, however, significant that today the presence of betalains in a family of angiosperms has become a key character used by all workers for its inclusion in the Centrospermae. Such families as the Cactaceae and Didiereaceae were allied with the other betalain 916 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 - clecvoge 0- cleavage HO b- cleavage HO HoN CO H # b- -cleovoge c ` — HOC ‘ | * ? COH CA HoN^ co COH HOC, NH; - " fa! =>. — N o“ "07 NO o `o“ con noc NN- coe Hoc H COH muscoflavin stizolobinic acid stizolobic acid betalamic acid (Amanita; Hygrocybe) (Mucong, Leguminosae) (Mucona, Leguminosae) etalains (Centrospermae; Amanita) GuRE 3. L-Dopa is known to undergo 2,3-1 (a-cleavage) or 4,5- (b-cleavage) extra diol cleavage in different organisms. Only in the Centrospermae and in mushrooms does L- opa lead via the b-cleavage pathway to betalamic acid, the precursor of all other betalains. I thank Prof. H. Musso, Univ. of Karlsruhe, for private discussions which led to the scheme presented here. families with a greater degree of confidence once their pigments were recognized. Of course, other data such as Jensen’s (1965) serological results also firmly es- tablished a close relationship of the Didiereaceae to the betalain families, espe- cially the Cactaceae and Portulacaceae. I suppose over the years the most controversial question has focused upon our 1963 (Mabry, Taylor & Turner, 1963) proposal that the Centrospermae be re- served for the betalain families, with the closely related but anthocyanin-produc- ing centrospermous families (e.g., Caryophyllaceae and Molluginaceae) being placed in a close but distinct order, the Caryophyllales. Although we did not then nor do we now concern ourselves with resolving the question of rank, the separation of the centrospermous families such as the Caryophyllaceae into a separate order “disturbed” a number of leading systematists. Therefore, our cur- rent treatment (Table 4), which still maintains distinct taxonomic categories (suborders) for the betalain and anthocyanin families, is more acceptable be- cause all the families are recognized as being “centrospermous.” While the phy- logenetic importance of the betalains is emphasized in our current treatment, the significance of other characters is recognized. It should be noted once more that our current treatment has been sharply influenced by the occurrence of the same P-III subtype sieve-element plastids in the Caryophyllaceae and Mollugina- ceae as are found in the betalain families. Despite having our views shaped by different kinds of data, the views of myself and many leading systematists are converging towards a common interpretation of the Centrospermae. 1977] MABRY—CENTROSPERMAE 917 TABLE 6. Pollen Morphology ( Nowicke, 1975). with Centrospermae-Specific Pollen: Some Taxa with Noncentrospermous Spinulose and Tubuliferous/Punctate Ektexine Pollen Betalain Families Achatocarpaceae Aizoaceae ataceae Amaranthaceae Gyrostemonaceae 5 Theligonaceae Cactacez Polygonaceae Chane sodas (incl. Dysphaniaceae ) Didiereaceae Nyctaginaceae Phytolaccaceae Portulacaceae Anthocyanin Families aryophyllaceae Molluginaceae A number of families sometimes treated as being centrospermous but which contain neither the P-III subtype sieve-element plastids nor betalains are now excluded from the Centrospermae proper (see Table 4, footnote a). Although the available molecular data do not suggest an alternative alignment for many of these taxa, this is not the case for the Bataceae and Gyrostemonaceae. - RNA hybridization data bear upon the relationship of the Bataceae to the Centro- spermae ( Chang & Mabry, 1974). First, these data indicate that the betalain fami- lies which were tested are closer to each other than to any other family and that the Caryophyllaceae is the closest family to the betalain group. At the same time, the Bataceae were clearly separated from the Centrospermae on the basis of the available DNA-RNA hybridization data. Moreover, it is significant that Prof. Martin G. Ettlinger, University of Copenhagen, recently detected (unpublished manuscript) benzylglucosinolate in Batis maritima L., a species previously re- ported to contain thioglucosidase (Schraudolf et al., 1971). On the basis of these and other data, Prof. Ettlinger allies the Bataceae with other glucosinolate- containing families (Ettlinger & Kjaer, 1968); furthermore, the isolation of an 5 from Codonocarpus cotinifolius (Desf.) F. Muell. (Bottomley & White, 1950) indicates that the Gyrostemonaceae also belongs with these same facile. As noted in the next section, the studies of pollen structure not only support a close relationship of Bataceae to the Gyrostemonaceae but also dis- tinguish these two families from those in the Centrospermae (Goldblatt et al., 1976; Nowicke, 1975 As a result of discussions with Prof. F. Ehrendorfer (Vienna) and from com- ments in a recent paper of his (1976), I agree that we must consider the possi- bility that the betalain families arose from an ancestral taxon which had lost the ability to produce anthocyanins. Such a process would require that the ances- tor had lost the one or two enzymatic steps required to convert dihydroflavo- nols into anthocyanins and subsequently gained the two or three steps needed to form betalamic acid from L-dopa and then condense this aldehyde with various amines and amino acids to produce the red and yellow betalains. 918 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 POLLEN MORPHOLOGY IN THE CENTROSPERMAE Recent investigations (Nowicke, 1975; Skvarla & Nowicke, 1976) of pollen morphology for centrospermous taxa (Table 6) also support a close relationship of the betalain families with the Caryophyllaceae and Molluginaceae in accord with our treatment (Table 4). In her examination of 190 species from 16 fami- lies by light and scanning electron microscopy, Nowicke (1975) found three common pollen types, all with a spinulose and tubuliferous/punctate ektexine among the betalain families and the Caryophyllaceae and Molluginaceae. Two additional minor related pollen types were detected in the Nyctaginaceae. The pollen morphology of other taxa such as Achatocarpaceae, Bataceae, Gyrostemonaceae, Polygonaceae, and Theligonaceae do not support their inclu- sion in the Centrospermae. Of these, only the Achatocarpaceae would appear to be centrospermous on the basis of having P-III subtype sieve-element plas- tids; the pigment content of this taxon is not yet known. SUMMARY It is our view that all the betalain families and the two anthocyanin families, the Caryophyllaceae and Molluginaceae, are derived from a common “centro- spermous” ancestral taxon which evolved the P-III subtype sieve-element plas- tids now characteristic of all these families. In addition, the ancestor was prob- ably preadapted for C, photosynthesis and for a pollen morphology with spinulose and tubiferous/ektexine. The ancestral taxon for the betalain families may have arisen either from a taxon which had anthocyanins, then lost them and later gained betalains, or from a taxon which had not previously contained either type of pigment. In any case, the centrospermous evolutionary complex is now repre- sented by eleven core families which in my opinion are best treated in one or- der, the Centrospermae or Caryophyllales, which consists of a betalain-suborder, the Chenopodiineae, and an anthocyanin-suborder, the Caryophyllineae (Table 4) LITERATURE CITED ARDENNE, R. H. Dörr, H. Musso & W. SrEGLicH. 1974. Isolation of muscaflavin from ra 7 5 species (Agaricales) and its dihydroazepine structure. Z. Naturf. 29C: 7-639. BEHNKE, H.-D. 1976a. Ultrastructure of sieve- element plastids in Caryophyllales (Centro— spermae), evidence for the delimitation and classification of the Order. Pl. Syst. Evol. 126: 4. ; Sieve-element plastids of Fouquieria, Frankenia (Tamaricales), and Rhab- dodendron "(Rutacede). taxa sometimes allied with Centrospermae (Caryophyllales). Taxon 25: 265-2 I $ed Transmission electron microscopy and systematics of flowering plants. Pl. Syst Evo T. J. Mn 1977. S-Type sieve-element plastids and uu in Viviania- ceae: evidence against ini inclusion into Centrospermae. Pl. Syst. Evol. 126: 371-375. * e „I. J. EIrERTr & T. J. Masry. 1974. Betalains se] P- 2 Pe sieve- tube plas- tids in N and 1 ( Phytolaccaceae). Taxon 23: 541-542 1 EY, W. & D. E. Warre. 1950. The chemistry of Western Australian plants, Part The essential oil of Codonocarpus cotinifolius ( Desf.) F. Muell. J. Proc. Roy. Austral. Chea. Inst. 17: 31-32. 1977] MABRY—CENTROSPERMAE 919 CuANG, C. & T. J. Masry. 1974. The constitution of the Order Centrospermae: RNA NA hybridization studies among betalain- and anthocyanin- producing families. Biochem. Syst. 1: 185-190. — ——-, L. Km™ter & T. J. Masry. 1974. Biogenesis of betalamic acid. Phytochemistry 13: 2771-2775. Cronguist, A. 1968. The Evolution and Classification of Flowering Plants. Houghton n. Dorp, H. & H. Musso. 1973. 5 Farbstoffe, II. . und Chromophore der Farbstoffe aus Amanita muscaria. Chem. Ber. 106; 3473-3482 1974. Chromatographic analysis of betalain pigments in toadstools and higher plants, Z. Naturf. 29C: Eckanpr, T. 1976. Classical Se n features of centrospermous families. Pl. Syst. Evol. 126: 5-25. EHRENDORFER, F. 1976. a remarks: systematics and evolution of centrospermous families. Pl. Syst. Evol. 126: 99-105. EICHLER, A. W. 1876. n. der Vorlesungen über Phanerogamenkunde. Schwerche IUE Kiel. —. Ru aap hea Zweiter Teil. W. Engelmann, Leipzig. ELLIS, B. i 1976. a ring-c iw in the biogenesis of spol is acid in Mucuna deeringiana. 5 15: 91. = M. G. & A. KJAER. Do Sulfur compounds in plants. In T. J. Mabry, R Iston & V. C. pnm a Recent Advances in Phytochemistry. Vol. 1; 59- 144. 1 Century-Crofts, N ork. FiscHER, N. & A. S. Sti. E Biosynthesis of betalains. On the cleavage of the aromatic ring during the enzymatic transformation of dopa into betalamic acid. Helv. Chim. à = 55: 649- E GOLDBLATT, W. NOWICKE J. Masry & H.-D. BEHNKE. 1976. Gyrostemonaceae: J. status bu affinity. Bot. Not. e 201-206. Hanporne, J. B. 1967. Comparative Biochemistry of the Flavonoids. Academic Press, ondon HUNZIKER, LH , H.-D. BEHNKE, I. J. Errerr & T. J. Masry. 1974. Halophytum ameghinoi: a lagala: contabiidé and Potype sieve-tube plastid species. Taxon 23: —539. IMPELLIZZERI, G. & M. PIATELLI. 1972. Biosynthesis of betalains: fo aon of indicaxan- thin in Opuntia ficus-indica fruits. Phytochemistry 11: 2499-2502. JENSEN, U. 1965. Serologische Untersuchungen zur Frage der systematischen Einordung er Didiereaceae. Bot. Jahrb. Syst. 84: 233-253. pus T. J. 1973. Is the Order Centrospermae monophyletic? Pp. 275-285, in G. Bendz & J. Santesson (editors), Chemistry in Botanical Classification. Nobel DU Lyon. Stock- holm and Academic Press, London 1976. Pigment dichotoniy and DNA-RNA hybridization data for centrospermous familias, Pl. Syst. Evol. 126: 79-94. & H.-D. BEHNKE (editors). 19764. Evolution of centrospermous families. Pl. Syst. me 126: 1-105. & H.-D. a 1976b. Betalains and P-type sieve-element 1 8057 The systema- tic position of e R. Br. (Centrospermae). Taxon 9-11 & A. S. . 1 The betalains. In T. J. Mabry, R. E. JM & V Runeckles (editors), p ee in Phytochemistry. Vol. 1: 145-160. Appleton- — Crofts, New York. — ———, H--D. “IFERT. 1976. Betalains and P-type sieve-element plastids in oh L. FFC Taxon 25: 112-114. „L. KI R & C. CHANG. 1972. The betalains: structure, function and biogenesis and the plant order Centrospermae. In V. C. Runeckles & T. C. Tso (editors), Recent Advances in Phytochemistry. Vol. 5: 104-134. Academic Press, New Yor . TAYLOR * & B. L. Turner. 1963. The betacyanins and their distribution. Phyto- chemistry 2: 61-6 „I. J. ne, CHANG, H. Masry, C. Kipp & H.-D. BEHNKE. 1975. Theligonaceae. Pigment d ultrastructural evidence which eee it from the order Centrospermae. Biochem. Syst. Ecc 3-55. JOWICKE, J. W. 1975. b morphology in the order Centrospermae. Grana 15: PrarrELLL, M. 1976. Betalains. In T. W. Goodwin (editor), Chemistry oat 1 of Plant Pigments. Vol. 1: 560—596. Academic Press, New York. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 220 Sarto, K., A. KOMAMINE & S. SENOH. 1975. Biosynthesis of stizolobinic i and stizolobic acid in the etiolated seedlings of Stizolobium rf. 30C: 659-662 1976. Further studies of the biosynthesis of stizolobinic ^i and stizolobic acid in etiolated seedlings of Stizolobium hassjoo. Z. Naturf. 3 URHBAUDQUN, „ B. SCHMIDT . WEBERLING, Das Vorkommen von 5 als Hinweis auf die 5 Stellung der Batidaceae. Experientia (Basel) 27: 1090- 109 SKVAR J. J. & J. W. Nowicxe. 1976. Ultrastructure of pollen exine in centrospermous families. Pl. Syst. Evol. 126: 55-78. TAKHTAJAN, A. Evolution und Ausbreitung der Blütenpflanzen. Gustav Fischer Verlag, Stuttgart. TIMBERLAKE, C. F. & P. BRIDLE. 1975. The anthocyanins. Pp. 214-266, in J. B. Harborne, abry & H. Mabry ag The Flavonoids. Chapman and Hall, London. WINTER, Evidence for the significance of crassulacean acid m as an adaptive mechanism to water stress. Pl. Sci. Lett. 3: 279-281 DEFENSIVE ECOLOGY OF THE CRUCIFERAE! PAUL FEENY? ABSTRACT The glucosinolates aniey oil glucosides), present in all crucifer species examined, seem to provide a major line of chemical defense against bacteria, fungi, insects, and mam- mals. 5 an suggests that other classes of secondary compounds, each re- stricted to one or a few genera, represent a second line of chemical defense Survival of wild crucifers depends partly on escape from 1 enemies in time and ided space. scovery of crucifers by several enemy species is aidec ee iw ei to glucosinolates or their breakdown is oducts. Pg med (ante rin eaves of Thlaspi arvense releases allylthiocyanate instead of the more typical allylisothiocyanate, which hi is used as a host-finding attractant by several insect species. E is change in secondary chemis- try may reduce the rate of 1 of Thlaspi plants by crucifer- adapted enemies. fensive ecology of crucifers seems to typify that of herbaceous plants generally: chemical resistance, in the n of small amounts of toxic compounds, combined with low apparency to enemies which are adapted to the chemical defenses. The importance of the Cruciferae and other families of herbaceous plants as sources of food-plants for man may result in large part from their relatively low concentrations of toxins. The mature foliage of trees, shrubs, and grasses, by contrast, remains poor food for man, just as for other plant enemies An a component of the defensive ecology of crucifers and other unapparent plants seems to be chemical diversity in space and time. Closer simulation of this diversity in fields of agricultural crops may reduce the need for synthetic pesticides The family Cruciferae comprises approximately 400 genera and 3,000 species, the vast majority of which are herbaceous (Vaughan et al., 1976). The greatest number of species are found in temperate regions of the northern hemisphere, especially in those with a Mediterranean type of climate. The Irano-Turanian region alone contains about 150 genera and 900 species and may well have been the . cradle of the inm at least in the Old Gg (Hedes, P regions sch some of the most dina 1 cible deserts, m it is poorly represented in the tropics ( Hedge, 1976). The family is the source of several economically important species and vari- eties, especially the cole crops of the genus Brassica. Economic incentives have stimulated extensive research on interactions between crucifers and their as- sociated insects and pathogens. Understanding of the chemical aspects of these interactions has been helped greatly by unusually thorough knowledge of the family's chemistry (see Kjaer, 1976). ! I thank the students in my general ecology class for compiling the list of human food- plants, Karen Arms for improving my grammar, and David Bates for considerable help with the preparation of the 5 Financial support was Bry vided by res search grant BMS- 7409868 from the National S ce Foundation and Hatch grant NYC-139413. epartment of 5 and Section of Ecology id S tonat Cornell University, Ithaca, New York 14853. ANN. Missouni Bor. Garp. 64: 221-234. 1977. 999 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 PRIMARY CHEMICAL DEFENSE The first characteristic line of chemical defense in crucifers is evidently pro- vided by the glucosinolates (mustard oil glucosides, thioglucosides). The oc- currence of these compounds, more than 70 of which are known, is restricted al- most entirely to the related families Capparaceae, Cruciferae, and Resedaceae. Approximately 300 crucifer species have been examined so far and all contain glucosinolates (Kjaer, 1960, 1976; Ettlinger & Kjaer, 1968). These compounds are hydrolyzed typically to yield volatile isothiocyanates (mustard oils) when plant tissues are damaged; allylglucosinolate (sinigrin), for example, is a major com- ponent of plants of the genus Brassica and breaks down to allylisothiocyanate: N-O-SO4* Z CH, = CH - CII -C + HO Ə CH» = CH- CH: - NCS + Glucose + HSO S-Glucose Allylisothiocyanate, released from allylglucosinolate, is largely responsible for the odor of cooked cabbage (MacLeod, 1976). Hydrolysis of glucosinolates is catalyzed by a group of enzymes (myrosinases) which are stored separately within the plant tissues but which come into contact with their substrates when the plant tissues are bruised or otherwise damaged (Kjaer, 1976; Bjórkman, 1976). Storage of isothiocyanates in the form of glucosinolates may represent an adaptation to avoid autotoxicity; isothiocyanates are strongly phytotoxic ( Hooker et al., 1945; Bell & Muller, 1973). Glucosinolates or their breakdown products are known to be powerful anti- biotics (Virtanen, 1958, 1965) and to inhibit the growth of fungi (Walker et al., 1937) and insects (Brown, 1951; Lichtenstein et al., 1964). The concentration of allylglucosinolate in the foliage of Brassica nigra plants in Tompkins County, New York, was found to be about 0.4% of fresh weight, depending somewhat on habitat and leaf age (P. Feeny and L. Contardo, in preparation); at this concen- tration the compound proved to be toxic to larvae of the black swallowtail but- terfly, Papilio polyxenes, which naturally feed on umbellifers but occur in the same habitats as many crucifer species in the northeastern United States ( Erick- son & Feeny, 1974; P. Blau, P. Feeny and L. Contardo, in preparation). Gluco- sinolates or their hydrolysis products, when ingested in large quantities, are also toxic to mammals; the effect seems to result, at least in part, from the effective- ness of allylisothiocyanate as a tissue irritant (Kingsbury, 1964). Glucosinolates in crucifers may play a role as allelopathic agents, inhibiting the germination and growth of competing plants. Patches within the annual grasslands of southern California are dominated by B. nigra, introduced from Europe. Bell & Muller (1973) showed convincingly that the persistence of these patches from year to year can be attributed to inhibition of the germination and growth of other plants by compounds leached from B. nigra. They found that allylisothiocyanate is a potent inhibitor of germination by seeds of several grasses but ruled it out as the allelopathic agent because of its rapid loss of activity in the soil. The unknown toxic compounds are water soluble and are leached from dead B. nigra tissues of the previous season's growth by the first fall rains, which 1977] FEENY—DEFENSIVE ECOLOGY 993 also serve as the stimulus for germination by the seeds of most species (Bell & Muller, 1973). I am not convinced that the authors have completely ruled out the possibility that the allelopathic agent is allylglucosinolate, stored in dead stems during the summer drought period and capable of releasing allylisothio- cyanate over a period of time after being leached into the soi The available evidence thus suggests that the biological activity of the glu- cosinolates is broad and supports the contention that predation, disease, and per- haps competition are selective pressures which have contributed to the evolution and diversification of these compounds in the Cruciferae (see also Feeny, 1976). THE CRUCIFER FAUNA In spite of their content of glucosinolates, crucifers are attacked by an ex- tensive array of insect species, several of which have become major pests of cul- tivated cruciferous crops. Many of the insect species which attack crucifers are “specialists” which rarely or never attack plants of other families; examples include larvae of the familiar cabbage butterfly, Pieris rapae, the cabbage aphid, Brevicoryne brassicae, and the cabbage flea beetles Phyllotreta cruciferae and P. striolata (Root, 1973). Generalist insects which include crucifers among their normal range of host-plants include the cabbage looper, Trichoplusia ni, and the green peach aphid, Myzus persicae. Crucifers are subject also to attack by an extensive array of fungi and bacteria (Westcott, 1971) and are probably eaten in significant quantities by wild mammals. In view of the deleterious effects of glucosinolates on organisms which do not normally attack crucifers we must presume that the various species making up the typical fauna of natural crucifers are somehow adapted to detoxify glucosinolates or otherwise avoid their harm- ful effects. The actual mechanisms of detoxification by adapted enemies are not yet known. Larval growth of the cabbage butterfly, P. rapae, on a wide range of crucifer species and cultivated varieties was compared recently by Slansky & Feeny (1977). Growth showed no obvious relationship to the varied pattern of glucosinolates present in the test plants but was closely related to the availability of nitrogen in the plant food. Individual glucosinolates vary in their toxicity to nonadapted insects (e.g. Brown, 1951); in crucifer-adapted insects, however, the extent to which tolerance of one glucosinolate confers tolerance of others needs to be examined in more detail. When concentrations of allylglucosinolate in the leaves of collard plants, Brassica oleracea, were artificially increased by culturing to 20 times the typical level, growth of P. rapae larvae remained unaffected (P. Blau, P. Feeny and L. Contardo, in preparation). This result suggests that glucosinolates represent “qualitative” or “evolutionary” barriers to nonadapted insects: once overcome by adaptation they have little or no toxic effect in spite of wide variation in con- centration (Feeny, 1975, 1976). This is consistent with the finding by van Em- den (1972) that relative growth rate of the cabbage aphid, B. brassicae, was correlated positively with the “total allylisothiocvanate" content of crucifer test- plants. Glucosinolates stimulate feeding by this crucifer-restricted aphid but they are evidently not toxic to it, at least at concentrations normally encountered in the plants. Dosage-related toxicity of glucosinolates may have greater ecologi- 994 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 cal effects on crucifer-adapted bacteria and fungi than it seems to have on in- sects which specialize on these plants. The effects of glucosinolates on generalist insects seem to be intermediate be- tween their effects on crucifer-specialists and those on insects which do not natu- rally attack crucifers. The southern armyworm, Spodoptera eridania, and peach aphid, M. persicae, naturally attack crucifers as well as plants of many other families. They must therefore be able to tolerate at least low levels of glucos- inolates. However, larvae of S. eridania have less tolerance for leaves artificially boosted with allylglucosinolate than have larvae of P. rapae (P. Blau, P. Feeny and L. Contardo, in preparation) and the relative growth rate of M. persicae is correlated negatively with the “total allylisothiocyanate" content of crucifer leaves ( van Emden, 1972). ESCAPE FROM ADAPTED ENEMIES IN SPACE AND TIME Such is the ability of adapted enemies to damage and destroy crucifer plants, once they have been discovered, that the survival of crucifers in nature must surely be attributable in large measure to their ephemeral life histories which provide a constantly changing pattern of geographical and phenological distribu- tion. The habitats most favored by crucifers seem to be those in which periods favorable for growth are severely limited by climatic variables such as rainfall (e.g. chaparral, grassland, desert) and temperature (arctic and alpine habitats). Typical crucifers must therefore be capable of rapid growth to maturity and seed-set, and it is perhaps not surprising that so many species have evidently been preadapted to exploit disturbed areas associated with human activities. Short growth season, shifting pattern of geographic distribution, and association with harsh and somewhat unpredictable climatic conditions are all characteristics which are likely to favor escape by plants from their adapted enemies (Janzen, 1970; Rhoades & Cates, 1976; Feeny, 1976). The importance of escape from adapted insect enemies to the ecology of herbaceous plants was well illustrated by the history of introduced Klamath weed, Hypericum perforatum, in California (Huffaker & Kennett, 1959). In 1951 this plant infested more than 2 million acres of range land, covering up to 80% of the ground area in some places. Introduction from Europe of the Hy- pericum-adapted leaf beetles, Chrysolina quadrigemina and C. hyperici, reduced the plant to less than 1% of its former abundance by 1959. Both plant and beetles continued to persist at low densities, the plant surviving best in shadier habitats where the beetles are less effective ( Huffaker & Kennett, 1959). "It would seem that the new low density of Hypericum perforatum is maintained at a level at which interplant distance restricts epidemic development of the beetle by limit- ing its opportunity to discover the isolated specific food plants" (Harper, 1969). While no such dramatic examples are available, it seems, for cruciferous plants, escape from discovery by adapted enemies is likely to be an important component of their defensive ecology also. Pimentel (1961) and Root (1973) have shown that populations of crucifer- adapted specialists such as B. brassicae and P. cruciferae reach higher densities on collard plants grown in monoculture patches than on plants grown among di- 1977] FEENY—DEFENSIVE ECOLOGY 995 verse meadow vegetation. Root (1973) attributed these findings to “resource concentration”: herbivores are more likely to find and remain on hosts that are growing in dense or nearly pure stands. An individual collard plant is more “ap- parent” (i.e., susceptible to discovery) when growing next to other collard plants than when growing among plants of other families ( Feeny, 1976). Comparable experiments by Smith (1976) showed that populations of B. Drassicae and other crucifer-feeding species reached higher levels on Brussels sprout plants grown on weed-free soil than on plants grown among weeds. Trapping experiments showed that weed-free plants were more attractive to colonizing aphids, prob- ably because a background of bare soil presents greater visual contrast than does a background of weeds (Smith, 1976). Diversity of surrounding vegetation may similarly permit wild crucifers in natural habitats to escape or reduce the risk of discovery by searching insects ( Feeny, 1976). Plants of the genus Dentaria differ from more typical crucifers in several re- spects. They are perennial and form patches, often of substantial area, among the ground vegetation of mature deciduous forests. The plants leaf out very early in the spring and approach senescence by the time the forest canopy has leafed out. Plants of D. diphylla were damaged heavily, after transplanting into open field habitats, by the typical open-habitat crucifer flea beetles P. cruciferae and P. striolata (Hicks & Tahvanainen, 1974) and Dentaria leaves supported better growth of P. rapae larvae than did those of any other crucifer tested (Slansky & Feeny, 1977). Though subject to their own specialized enemies, such as the butterfly Pieris virginiensis and the flea beetle Phyllotreta bipustu- lata, Dentaria species have probably benefited by their escape, in evolutionary time, into a habitat which is atypical of crucifers and thus not frequented by many of the typical crucifer-adapted enemies. PLANT-FINDING ADAPTATIONS Many crucifer-adapted insects have evolved behavioral responses to glucosi- nolates or their breakdown products, thus permitting them to find their food- plants more easily and to discriminate them from other vegetation. An early example of such behavior was described by Verschaffelt (1911) who found that larvae of P. brassicae and P. rapae can be stimulated to feed on normally re- jected plants by treating the plants with solutions of allylglucosinolate. A recent review by Schoonhoven (1972) lists a dozen insect species which are known to make use of these compounds as behavioral cues. There is even a crucifer- adapted fungus, Plasmodiophora brassicae, the spores of which are stimulated to germinate by the presence of allylisothiocyanate (Hooker et al, 1945). Be- havioral responses to glucosinolates or isothiocyanates by individuals of any one insect species usually depend on concentration and may also vary from one compound to another (e.g. Thorsteinson, 1953; Hicks, 1974; Finch & Skinner, 1974) The crucifer-feeding flea beetles, Phyllotreta cruciferae and P. striolata, are strongly attracted to traps containing solutions of allylisothiocyanate (Feeny et al., 1970) and can also be induced to eat bean leaves, which they normally re- ject, when these have been cultured in solutions of allyglucosinolate (Hicks, 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 1974). We have found recently that addition of vials containing solutions of allylglucosinolate in mineral oil to 3-plant islands of Brassica nigra, planted among diverse vegetation, greatly accelerated the rate of discovery of the plants by these flea beetles (P. Feeny, J. Gaasch and L. Contardo, in preparation). This finding not only confirms the effectiveness of allylisothiocyanate as a host-find- ing attractant but also shows that leakage of such compounds, even in small amounts, can be a liability to B. nigra plants since it increases their apparency to adapted enemies. SECONDARY DEFENSE IN CRUCIFERS Many crucifers are known to contain other secondary compounds in addition to glucosinolates. The genera Erysimum and Cheiranthus, for example, contain cardenolides, the genus Iberis contains cucurbitacins, and plants of the genera Lunaria and Capsella contain alkaloids ( Gheorghiu et al., 1959; Hegnauer, 1964). The genera Lepidium and Thlaspi contain atypical enzymes which break down glucosinolates not to the typical isothiocyanates but to their 88 geo- metrical isomers, the thiocyanates (Gmelin & Virtanen, 19 Many of these plants are avoided by crucifer- adapted insects or, if fed upon, support unusually poor growth. Larvae of P. rapae, for instance, grow poorly on Thlaspi arvense, Lepidium virginicum, and Lunaria annua (Slansky & Feeny, 1977); they will refuse to eat leaves of Erysimum cheiranthoides and Capsella bursa-pastoris (A. M. Shapiro, personal communication). Verschaffelt (1911) found that C. bursa-pastoris was attacked only very slightly by larvae of P. rapae and P. brassicae; E. perofskianum was also less preferred by these larvae relative to most other crucifers offered to them. Plants of E. cheiranthoides, C. bursa- pastoris, and Iberis amara are not eaten by P. cruciferae flea beetles (Feeny et al., 1970). Chew (1975) found that larvae of Pieris napi macdunnoughii in Colorado refused to eat Erysimum asperum. Larvae of P. napi macdunnoughii grew normally on Thlaspi montanum, a native plant in Colorado, but they and larvae of P. occidentalis died after eating T. arvense, an introduced species. The unusual resistance of plants of these genera to typical crucifer enemies may re- sult from their content of atypical secondary compounds (see Verschaffelt, 1911). Allylthiocyanate, for example, is known to be toxic to insects (Brown, 1951). Such compounds could have been evolved as a "second line of defense" in re- sponse to enemies which have evolved mechanisms for tolerating glucosinolates and their typical hydrolysis products. Diversification of secondary chemistry, in other words, may permit escape from certain enemies in evolutionary time, at least until further counteradaptations are evolved by the associated insects or other enemies. In addition to their possible toxic or growth-inhibitory effects, unusual sec- ondary compounds may further benefit a plant species by reducing apparency to adapted enemies. Hydrolysis of allylglucosinolate in leaves of Thlaspi arvense yields allylthiocyanate instead of the more typical allylisothiocyanate (Gmelin & Virtanen, 1959; P. Feeny and L. Contardo, in preparation). Three-plant is- lands of T. arvense were colonized by Phyllotreta flea beetles at a considerably slower rate than were nearby islands of Brassica nigra, perhaps because allyliso- 1977] FEENY—DEFENSIVE ECOLOGY 997 - thiocyanate is an attractant to the beetles whereas allylthiocyanate is not. Coloni- zation of T. arvense islands was accelerated by addition of vials containing solu- tions of allylisothiocyanate (P. Feeny, J. Gaasch and L. Contardo, in preparation ). Crucifers may derive additional protection from adapted enemies as a result of association with plants of different chemistry. Tahvanainen & Root (1972) have found that odors from tomato, Lycopersicon lycopersicum (=esculentum), and ragweed, Ambrosia artemisifolia, plants interfered with the ability o cruciferae flea beetles to find crucifer host-plants. The reduction of plant ap- parency to enemies by neighboring plants of different species is an important component of “associational resistance" (Tahvanainen & Root, 1972)—a2 phe- nomenon frequently exploited by organic gardeners. CHEMICAL DEFENSE AND THE HUMAN DIET The defensive ecology of crucifers seems to typify that of many ephemeral herbaceous plants—plants which rely to a great extent on being hard to find (unapparent) in natural habitats. Such plants seem to contain rather low con- centrations of effective toxins. They probably benefit from a diversity of chemi- cal defense in any one species and from association with other plants of different chemistry ( Rhoades & Cates, 1976; Feeny, 1976). Their defenses clearly differ from those of the mature foliage of more persistent plants such as shrubs and trees. Such plants are bound to be found by enemies and must correspondingly be well defended; they often contain large amounts of general growth-inhibitory compounds, like tannins, resins and silica, which are resistant to simple counter- adaptation. The foliage of apparent plants is usually tough and deficient in nutrients and water when compared with that of most herbaceous plants ( Rhoades & Cates, 1976; Feeny, 1976) These differences in the defensive ecology of plants, depending upon their apparency to enemies, seem to be reflected in human food preferences. One hundred students in the general ecology course at Cornell University were asked to list as many species of human food-plants as they could think of in 15 minutes. Their total of 108 species, excluding plants used primarily as spices and drugs, was then tabulated by plant growth form and by what part of the plant is eaten (Fig. 1 and Appendix). Though undoubtedly a biased view of more general patterns of plant consumption by man, this survey revealed some interesting and suggestive trends. Most of the species listed are harvested only for their fruits or seeds, and of these species most are trees (Fig. 1). The production of fleshy fruits is probably an adaptation for seed dispersal by vertebrate animals, including our primate ancestors. Ripe fruits are adapted to be attractive to animals by their size, color, and taste; fruit-eating behavior by the animals is probably reinforced by the fact that fruits contain not only energy-rich carbohydrates and fats but also vita- mins and mineral ions which are vital for the survival of many vertebrate ani- mals and not readily available from other natural sources (see McKey, 1975). One can even speculate that our "sweet tooth," now a conspicuous liability in times of readily available sugar, represents a physiological adaptation which 998 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Tree 38 Shrub or perennial vine 4 Herb 16 Grass Root Foliage Fruit / Seed RE l. Distribution of 108 species of human food-plants according to plant growth form ind part of plant eaten. Figures d u. of food-plants listed in each category (3 species listed twice). See Appendix for deta stimulated our ancestors to seek out fruits with their high nutrient value (see Yudkin, 1969). A second striking pattern reflected in this survey is that plants whose roots or leaves form part of the human diet are almost all herbs (Fig. 1). These are the plants, including the ancestors of our cruciferous vegetables, which tend to be ephemeral and unapparent in nature. The origins of cultivated plants from herbaceous species were probably due to the concentrated food value of the roots or tubers of many of these species and to the unique preadaptations of 1977] FEENY—DEFENSIVE ECOLOGY 999 “weedy” plants to thrive in the disturbed habitats associated with human habita- tion (Hawkes, 1969). Preferential consumption of herbaceous species may also reflect the presence in trees and shrubs of extensive chemical and physical de- fenses, evolved by trees and shrubs because they are relatively apparent to natural enemies. Many of the drugs and spices used by man come from the foliage and roots of trees and shrubs, though they are rarely consumed in large quantities. By contrast we are presumably able to tolerate the comparatively low concen- trations of defensive compounds in crucifers and other herbaceous plants both because of the detoxication enzymes concentrated in the vertebrate liver ( Free- land & Janzen, 1974) and also, since the cultural evolution of the use of fire, be- cause we further detoxify or remove many of these compounds by cooking (Yud- kin, 1969; Leopold & Ardrey, 1972). Only because they contain relatively small concentrations of toxins can we consume such plants in large quantities. APPARENCY AND AGRICULTURE The effectiveness of natural plant defenses is reduced by present agricultural methods. When they are planted in monocultures, crop plants become more apparent to natural enemies than are their ancestors in nature, yet they possess chemical and physical defenses inappropriate for survival as apparent plants. This is a major reason that substantial quantities of synthetic pesticides are often required to prevent widespread devastation of crops. It would undoubtedly be possible to modify crop varieties and agricultural methods so as to mimic the defensive ecology of wild ancestral plants more closely. Levels of natural defensive compounds could be maintained or restored by selective plant breeding and emphasis placed on diversity of defense within any particular crop species. Plant apparency could be reduced by such tradi- tional techniques as crop rotation and interplanting of different crops or chemi- cal varieties of any one crop. Apparency could be reduced further by eliminating or modifying those plant chemicals which the more important plant enemies use as behavioral attractants or feeding stimulants. Strategies to improve and diversify chemical resistance would be more ef- fective if they were coordinated with strategies to reduce plant apparency. Just as the evolution of resistance to a particular pesticide by an insect population may result from extensive and repeated exposure to that compound, so also the ewer the insects which find a particular plant variety, the less likely they are to evolve methods of tolerating the plant’s chemical defenses (Southwood, 1973). A key component of the defensive ecology of crucifers and other unapparent plants seems to be chemical diversity in space and time (Rhoades & Cates, 1976; Futuyma, 1976; Feeny, 1976). The more closely we can simulate this diversity in our fields of vegetable crops, the less dependent are we likely to become on the use of synthetic pesticides to achieve a given level of agricultural production. LITERATURE CITED BELL, D. T. & C. H. Murer. 1973. Dominance of California annual grasslands by Bras- sica nigra. Amer. Midl. Naturalist 90: 277—299. Björkman, R. 1976. Properties and function of plant myrosinases. Pp. 191-205, in 930 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 J. G. Vaughan, A. J. MacLeod & B. M. G. Jones (editors), The Biology and Chemistry of the Cruciferae. Academic Press, New Yor Brown, A. W. A. 1951. Insect Control by Chem icals. Wiley, New York. Cuew, F. S. 1975. Coevolution of pierid butterflies and their cruciferous foodplants. I. The relative quality of available resources. Oecologia 20: 117-127. ERICKSON, J. P. FggNy. 1974. Sinigrin: A chemical barrier to the black swallowtail butterfly, Papilio is Soni rn 55: 103-111. ETTLINGER, A. Kjaer. 1968. Sulfur ka s; in plants. In T. J. Mabry (edi- b. cd. uos in Boe. Vo FEENVY, P. 1975. Biochemical coevolution . plants and their insect herbivores. Pp. 3-19, in L. E. Gilbert & "u H. Raven (editors), cuu of Animals and Plants. Univ. of Texas Press, Aust 976. Plant 5 and chemical defense. ja J. W. Wallace & R. L. Mansell ( editors), Recent Advances in ET ud Mos 1-40. . L. PAAUWE & EMONG. d tles and mustard oils: Host plan specificity of Phyllotreta cruciferae i P. 0 adults (Coleoptera: 5 Ann. Entomol. Soc. Amer. 63: 832-841 Fincu, S. & G. SKINNER. 1974. Studies on the cabbage root fly: Evaluation "i attractants. Rep. Natl. Veg. Res. Stat. ( Wellesbourne, Warwickshire, England) for 197 84—85. FREELAND, W. J. & D. H. Janzen. 1974. Strategies in herbivory by aedi the role of plant p a eas Amer. Naturalist 108: 269-289. Furuyma, D. Food plant specialization and environmental predictability in Lepi do optera. ds Naturalist 110 285-2 id GHEORGHIU, A., CONSTA & E. Ionescu-Matiu. 1959. Extragerea si separarea alcaloizilor fluorescenti din sits bursa nes L. Stud. Cercet. Biochim. 2: GMELIN, R. & A. I. VIRTANEN. 1959. A new type of m cleavage of mustard oil glucosides. Formation of allylthiocyanate in Thlaspi arvense L. and benzylthiocyanate in Lepidium dcin L. and Lepidium sativum L. sks Chem. Scand. 13: 1474-1475. PAP LL. role of predation in vegetational diversity. In G. M. Woodwell H. H. ros (edito) Diversity od St tability | in ss sisal] Systems. Brookhaven Symposium in Biology. N 62. Hawkes, J. 1969. The fae ical background of plant domestication. Pp. 17-29, in P. J. D G. W. Dimbleby (editors), The Domestication and Exploitation of Plants and 1 Duckworth, London. * I. C. 1976. A systematic and 5 survey of the Old World Cruciferae. 1-45, in J. G. Vaughan, A. J. MacLeod & B. M. G. Jones (editors), The Biology and 8 of the Cruciferae. Academie Press, Wew ork. HEGNAUER, R. 1964. SEEE N der Pflanzen. Band 3: Dicotyledoneae: Acantha- eae—C US Berkhäuser Verlag, Basel. Hicks. K. 4. Mustar | oi glucosides: feeding stimulants for adult cabbage flea beetles, Phyllotreta cruciferae (Coleoptera: Chrysomelidae). Ann. Entomol. Soc. Amer. 264 & J. O. TAH VANAINEN. 1974. Niche differentiation by 3 . flea beetles (Coleoptera: C E lial Amer. Midl. Naturalist 91 Hooker, W. J., WALKER & K. P. Linx. 1945. Effects of aoe oils on Plas- modiophora brassicae and their relation to resistance to clubroot. J. Agric. Res. 70: 63-78. HurFaker, C. B. & C. E. KENNETT. 1959. A ten year study of the vegetational changes as ssociated with biological control of Klamath weed. J. Range Managem. 12: 69-82. Janzen, D. H. 1970. Herbivores and the 1 of tree species in tropical forests. Amer. Naturalist i d 501-528. Kincspury, J. M. 1964. pen Plants of the United States and Canada. Prentice-Hall, Inc., Englewood Cliffs, New Jer KJAER, A. 1960. Naturally derived isothiocyanates (mustard oils) and their parent gluco- sides. Fortschr. Chem. Org. Naturstoffe 18: 122-176. 1976. rn in the Cruciferae. Pp. 207-219, in J. G. Vaughan, A. J. M Leod B . M. G. Jones (editors), The Biology and Chemistry of the Cruciferae. read demic Press, New York. LEoPoLp, A. C. & R. ARpREY. 1972. Toxic substances in plants and the food habits of early n : LICHTENSTEIN, E. P., D. C. Morcan & C. H. MUELLER. 1964. Naturally occurring insecti- cides in cruciferous crops. J. Agric. Food Chem. 12: 158-161. 1977] FEENY—DEFENSIVE ECOLOGY 23] MacLkEop, xa ] 1976. E flavour compounds of the Cruciferae. Pp. 307— in . G. Vaughan, A. J. MacLeod & B. M. os Jones (editors), The Biology and Lesa k of the Cruciferae. e Press, New "ms D. 1975. The ecology of iud seed dispersal systems. Pp. 159-191, in L. ilbert & P. H. Raven (editors), Coevolution of Animals and Plants. Univ. of Texas 55 Austin. * D. 1961. Species diversity and insect population outbreaks. Ann. Entomol. Soc. Amer. 54: f Ruoaves, D. F. C. Cates. 1976. Toward a general theory of plant antiherbivore chemistry. m J. Ea Wallace & R. L. Mansell (editors), Recent Advances in Phytochemis- try. Vol. 10: 168-213 Roor, R. B. 1973. Organization of a plant-arthropod association in simple and diverse habitats: the fauna of collards (Brassica oleracea). Ecol. Monogr, 43; 95-124. SCHOONHOVEN, L. M. 1972. Secondary plant substances and pn. s + V. C. Runeckles & T. C. Tso (editors), Recent Advances in Phytochemistry. Vol. 197—224. SLANSKY, F., In. & P. FggNY. 1977. Stabilization of the rate of nit E accumulation by in of the cabbage butterfly on wild and cultivated food-plants. Ecol. Monogr. 47: 209-22 SMITH, J. G. 1976. Influence of crop ge on aphids and other phytophagous in- sects on Brussels sprouts. Ann. Appl. Biol. -13. SOUTHWOOD, 5 R. E. 1973. The insect ds 1 evolutionary perspective. Pp. 3-30, in H. F. van Emden (editor), Insect, Plant Relationships. Blackwell Scientific Publications. Oxford. TAHVANAINEN, & R. B. Roor. 1972. The influence of vegetational diversity on the population ae of a 5 herbivore, Phyllotreta cruciferae (Coleoptera: Chrysomelidae ). es 10: 321-346. TuonsrEINSON, A. J. 1953. The chemotactic responses that determine host specificity in olig gophagous insect (Plutella maculipennis (Curt.) Lepidoptera). Canad. J. Zool. : 2- 72, VAN EMDEN „H. F. 1972. Aphids as phytochemists. Pp. 25-43, in J. B. Harborne ape rhe Ecology. Academic Press, n on. VAUGHAN, J. G., A. J. MacLEop & B. M. JONES (editors ). 1976. The Biology and Chemistry of the E Academic "dion New Yor VERSHAFFELT, E. 191 The cause determining the letina of food in some herbivorous insects. Proc. he Sei. Amsterdam 13: 536-542. VIRTANEN, A. I. 1958. Antimikrobiell wirksame substanzen in kulturpflanzen. Angew. 52. — editor), ——— 965. Studies on p sulphur compounds and other labile substances in plants. Phytochemistry 4: -228. WALKER, J. C ORELL & H. H. Foster. 1937. í of mustard oils and related sul- — Feompennds S certain fungi. Amer. J. Bot. 24: 536-54 EST „C. 1971. Plant Disease Handbook. Van aa es ae New York. ae n 1969. 1 and the nutritionist. Pp. 547-552, in J. Ucko & G. W. — gn i The Domestication and Exploitation of Plants pu Poss Duck- orth, London 939 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 APPENDIX Species of human food-plants listed by 100 students in the general ecology course (Bio. Sci. 360, Fall 1976) at Cornell University, and categorized as a function of: (1) Growth form of plant and (2) Part of plant eaten. Bulbs and tubers are included with roots; shoots, stems and flower parts are included with foliage. Three species Vitis vinifera, ~ vulgaris, and Brassica rapa) are listed twice. an. = annual, bien. = biennial, per. = perennial. A. TREES (i) Root: No species listed. (ii) ursi No species listed. ) Fru (iii Mango Mangifera indica tree Anacardiaceae Pawpaw Asimina triloba small tree Annonaceae Papaya Carica papaya small tree Caricaceae Japanese persimmon Diospyros kaki tree Ebenaceae iin, ado 5 americana tree Lauraceae Fic tree oraceae Breadfruit 1 ‘altilis tree Moraceae Banana, plantain Musa acuminata and tall per. herb Musaceae x paradisiaca tall per. herb Musaceae Common guava Psidium guajava small tree M Olive lea europaea ree Oleaceae ate Phoenix dactylifera tall palm Palmaceae Pomegranate Punica granatum small tree Punicaceae uince Cydonia oblonga small tree Rosaceae pple Malus pumila tree Rosaceae Pear Pyrus communis tree Rosaceae ricot Prunus armenica small tree Rosaceae Sweet cherry Prunus avium tree Rosaceae Plum Prunus domestica small tree Rosaceae Peach, — Prunus persica small tree Rosaceae Sweet orange Citrus sinensis tree Rutaceae Grapefru rit Citrus x paradisi small tree Rutaceae Nagami kumquat Fortunella margarita small tree Rutaceae (iv) Seed Cashew Anacardium occidentale tree Anacardiaceae istachio Pistacia vera small tree Anacardiaceae European filbert/hazelnut Corylus avellana small tree Corylaceae American d eM Corylus americana small tree Corylaceae European ches C i tree aga Beech Fagus grandifolia tree Fagaceae ecan Carya illinoensis tree Juglandaceae Hickory Carya ovata an tree Juglandaceae Carya lacinio tree Juglandaceae Butter Juglans cinerea tree Juglandaceae English dant Juglans regia tree Juglandaceae Brazil nu Bertholletia excelsa tree Lecythidaceae i. um Cocos nucifera tall palm Palmaceae Pinus cembroides ree Pinaceae 1 Prunus dulcis small tree Rosaceae B. SHRUBS AND PERENNIAL VINES (i) Root: Sweet potato (tuber) Ipomoea batatas per. vine C 8 vam Dioscorea spp. per. vine Diosco Cassava/manioc Manihot esculenta shrub Euphorbiaceae Ground nut (tuber) Apios americana per. vine Leguminosae (ii) Foliage: European grape Vitis vinifera per. vine Vitaceae 1977] FEENY—DEFENSIVE ECOLOGY 933 APPENDIX. (continued) (ii) Fruit: American elder Sambucus canadensis shrub Caprifoliaceae Huckleberry Gaylussacia sp shrub Ericaceé Cranberry 14 macrocarpon shrub Ericaceae Blueberry Vaccinium sp shrub Ericac Passion fruit/purple Passiflora edulis per. vine Passifloraceae nadill Red raspberry Rubus idaeus shrub Rosaceae Black raspberry Rubus occidentalis shrub Rosaceae Loganberry, boy senberry Rubus ursinus shrub Rosaceae Rose hi Ros shrub Rosaceae American gooseberry Ribes pou shrub Saxifragaceae Red currant Ribes sativum shrub Saxifragaceae Fox grape Vitis labrusca per. vine Vitaceae Mu e grape Vitis vinifera per. vine Vitaceae Seed: No species listed. C. HERBACEOUS PLANTS (i) Root Onion (bulb) Allium cepa per. Amaryllidaceae Beet, sugar beet Beta vulgaris an./bien Chenopodiaceae Rutabaga Brassica napus an./bien. "ruciferae Turni Brassica rapa /bien Cruciferae Radish Raphanus sativus an.) bien. Cruciferae Burdock Arctium lappa per. Compositae Jerusalem artichoke (tuber) Helianthus tuberosus per. Compositae Camass (bulb) amassia quamash per. Liliaceae Potato (tuber) Solanum tuberosum per. Solanaceae Cattail Tupha spp. per. Typhaceae Carrot Daucus carota an./bien. Umbelliferae Parsnip Pastinaca sativa bien Umbelliferae 0 Foliage: Leek Allium ampeloprasum bien. Amaryllidaceae n) Symphytum officinale Der. Boraginaceae eet Be ilgaris an. / bien. Chenopodiaceae pin: "i Spinacea oleracea an. / bien. . x soap kale, ete. Brassica oleracea an./bien. Crucifera Cine s. ‘cabbage Brassica rapa an./bien. 8 ater cress Nasturtiu Nid tga per. Cruciferae mu vont end, an./bien. Compositae Chic Cichorium 0 5 per. Compositae pee (flower bud and Cynara scolymus per. Compositae scales ) Lettuce Lactuca sativa an./ bien. Compositae Dandelion Taraxacum officinale per. C 8 sparagus (young stem) Asparagus officinalis per. Liliace Rhubarb (leaf stalk) Rheum rhabarbarum per. 1 ae Celery (leaf stalk) Apium graveolens bien. Umbelliferae Fennel (leaf stalk ) Foeniculum yas in an./per. Umbelliferae (ii) Fruit: Pineapp Ananas comosus per. Bromeliaceae Sunflower Helianthus annuus an. Composit Watermelon Citrullus lanatus an. vine Cucurbitaceae Melon Cucumis melo an. vine Cucurbitaceae Cucumber Cucumis sativus an. vine Cucurbitaceae Squash, EOS, zucchini Cucurbita s an. vine Cucurbitaceae Strawber ragaria x ananassa per. Rosaceae Green m chili Capsicum annuum an./per. Solanaceae 934 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 APPENDIX. (continued) Tomato Lycopersicon Vina: an./per. Solanaceae Eggplant olanum melongen an./per. Solanaceae (iv) Seed: Peanut Arachis bri an. Leguminosae Soybean Glycine m an. Leguminosae enti Lens cul n. Leguminosae Lima bean Phaseolus limensis an./per. Leguminosae Kidney bean Phaseolus vulgaris an i Garden pea Pisum sativum an. vine Leguminosae D. GRASSES (i) Root: No species listed. (ii) Foliage: Bamboo (young shoots ) Phyllostachys spp. and per. 5 mbusa spp. per. Gram Sugar cane (stems Saccharum officinarum per. Coe iii) Fruit: No species listed. (iv) I Oat Avena sativa an. Gramineae Mro Hordeum vulgare an. Gramineae ce Oryza sativa an. Gramineae AEA /millet Panicum miliaceum an. Gramineae Rye Secale cereale an. Gramineae Sor hu Sorghum bicolor an. Gramineae rgnt 8 Wheat Triticum aestivum an. Gramineae Corn an. Gramineae CHEMOSYSTEMATICS AND ITS EFFECT UPON THE TRADITIONALIST' B. L. TURNER? What is a traditionalist, taxonomically speaking? I suppose a traditionalist might best be defined as a taxonomist trained as a pheneticist, practicing his trade as a pheneticist, and constructing his classification using primarily phenetic data. By this definition I am a traditionalist and consequently can claim to answer, for myself, the effect of chemosystematics upon my own traditional at- titudes and outlooks. And this has been profound. I say profound not because this new field has solved any large number of critical problems in plant taxonomy, but because where it has been used with skill and judgement, it has proved much more effective than phenetics in solving the particular problems concerned. Indeed, without chemical data many of the more intractable problems having to do with familial relationships among flower- ing plants generally are likely to remain unresolved: there are simply too many cooks and nearly all with varying tastes. Even if they all see the same phenetic substances in the phyletic cabinet, they nonetheless are prone to come up with different combinations of this or that ingredient (selected characters), with varying amounts (intuitive weighting), to say nothing of the condition (basic I. O.) or temperature (zealousness) of the oven (i.e. brain). I suspect that most traditionalists, even some of the best, do not like to be reminded that their approach is fraught with such variables, or that data de- rived from some other discipline might prove superior to those from their own. As an example, when the late Dr. Alston and I first showed the utility of paper chromatography for resolving problems of natural hybridization in Bap- tisia, an eminent, not so classical, plant systematist suggested that our documen- tation of complex hybridization in this genus could have been accomplished with equal clarity using selected morphological characters arranged upon Anderson- type scatter diagrams. Needless to say this intellectual guffaw was issued by the late Edgar Anderson, and the ironic part of all this is that Anderson himself was the first to collect and call attention to the existence of hybrid swarms among this group of plants ( Anderson, in Larisey, 1940b), but he failed to perceive its complexity, in spite of the fact that he collected his hybrid populations of Bap- tisia in a region where the potential for trihybridization is not infrequent (Alston & Turner, 1963). In fact, I seriously doubt that Anderson, or any traditional sys- tematist, including myself, would have been able to recognize, much less intuit, trihybridization within this group, to say nothing of its documentation with rea- sonable certainty using morphological characters. Trihybridization, of course, is rather the exception in nature: most species tend to comingle two at a time at any one site. But even then, lacking in situ clues (for example, two parental taxa occurring together with their putative isopod in part, by NSF Grant DEB 76-09320. > Botany Department, The University of Texas, Austin, Texas 78712. ANN. Missouni Bor. Garp. 64: 235-249. 1977. 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 hybrids, as happens with Baptisia upon occasion), two-way hybrids may be difficult to detect, especially where these are quite distinct and relatively widely distributed. Hence we find the southeastern taxon, Baptisia serenae Curtis, be- ing recognized as a good species for over 195 years by a wide range of workers, including such an outstanding traditional worker as Wilbur (1963). It is, how- ever, an F, hybrid of B. tinctoria (L.) Vent. x B. alba (L.) Vent. A more re- markable F, hybrid, also long-recognized as a good species by nearly all tradition- alists, including the only recent monographer of the genus (Larisey, 1940a) is B. microphylla Nutt., this being the relatively rare hybrid between B. perfoliata (L.) R. Br. and B. lanceolata (Walt.) Ell. While it is perhaps likely that these very distinct F, hybrids would have been detected if they were found growing with their putative parents, reasonable verification, short of long and laborious crossing experiments, would be difficult. If, however, their flavonoid profiles were sufficiently different, putative F, hybrids might be readily confirmed, even from hybrids mounted on herbarium sheets up to 100 years old. Speaking of the superiority of micromolecular data for the resolution of sys- tematic problems, the most telling example of its efficacy is that involving the detection of allopatric introgression. Edgar Anderson (1949) in an Epilogue to his brilliantly conceived text, Introgressive Hybridization, made the following statement: How important is introgressive hybridization? I do not know. One point seems fairly certain: its importance is paradoxical. The more 5 introgression becomes, the greater is its biological significance. It may be of the greatest fundamental importance when by our present crude methods we can do no more than to demonstrate its existence. en, the other hand, it leads to bizarre hybrid swarms, apparent even to the casual passer-by, it may be of little general 5 cx. Only by the exact comparisons of populations can we demonstrate the phenomenon . . e irs spread of a few genes (if it exists) might well be 5 even from a study of Sor daten averages, but it would be of tremen- dous biological import . . . . Hence our paradox ind d ession is of the greater biological sig- nificance, the less is the impact apparent to casual inspecti In other words, in well-differentiated, sympatric species such as Baptisia where natural hybridization can be easily recognized and readily documented, its biological impact on evolutionary processes is negligible. But in allopatric situations where hybridization is very difficult to detect it is likely to be of the greatest biological significance. In spite of these reflections from the foremost proponent of introgressive hybridization, few, if any, well-documented studies have been forthcoming on allopatric introgression. In fact, the best documented case in the literature for allopatric introgression is reportedly that involving Juniperus virginiana L. and J. ashei Buchh. ( Anderson, 1953; Davis & Heywood, 1963). However, in a num- ber of detailed studies, centered at The University of Texas (using 60 to 80 chem- ical characters as detected by gas chromatography as well as morphological characters which distinguish between the taxa), the existence of F, hybrids or their immediate derivatives could not be detected, even at sites where large populations of both species grew intermixed, and in no instance could the exis- tence of introgression be inferred from the data accumulated (Flake, von Rud- loff & Turner, 1969; Adams & Turner, 1970; Flake, von Rudloff & Turner, 1973). In short, what was taken to be a very well-documented case study of allopatric 1977] TURNER—CHEMOSYSTEMATICS AND TRADITIONALISTS 937 introgression turned out to be a situation in which clinal intergradation in habital features over a broad region occurred such that, superficially, hybridization and introgression might be inferred. In hindsight, it now seems rather reasonable to have viewed the case study of introgression between J. virginiana and J. ashei with considerable doubt, for the two species are readily distinguished by a number of morphological features and are placed in different species—groups (sections) of the genus, and the character used for such taxonomic segregation (cilia along the leaf margins) does not, to our knowledge, segregate in putative hybrid swarms (ie., the two species can always be recognized by this feature, and others, as attested to by the repeated correlation of this character with a plethora of chemical characters); other experienced field workers such as D. S. Correll (pers. comm.) have also had no difficulty in placing the plants concerned in one taxon or the other. In fact, as already indicated, the morphological variation found in J. virginiana is clinal, i.e., the species has formed or is in the process of forming regional races as a result of adaptational mechanisms arising out of its own gene pool, this be- ing unrelated to the possible influx of genes from the largely allopatric J. ashei. Our work has substantiated fully these suppositions (Flake & Turner, 1973). Again, it is ironic that Hall, who was Anderson's student, should have docu- mented introgressive hybridization where this was not occurring. We attribute this to the plasticity of the morphological characters selected for its detection. It was the absoluteness of the chemical data themselves which permitted reso- lution of the problem. What we were left with then was no well-documented case study of al- lopatric introgression of a regional nature. Fortunately, however, there has been a recent, carefully conceived, populational study of J. virginiana and J. scopulorum Sarg. in the Missouri River Basin of the north central United States by Van Haverbeke (1968a) which appears to be a situation involving allopatric intro- gression of the type Anderson felt to be so important in evolutionary processes. The study seems to be unusually well documented. Van Haverbeke made very accurate records of the populational sites, including precise data on ten indi- vidually marked trees which were selected for study at each site. These included photographs and detailed field notes. In short, the J. virginiana-]. scopulorum complex appeared to provide an ideal case study of allopatric introgression using the chemonumerical methods that proved so effective in disproving the oc- currence of this phenomenon in the J. ashei-J. virginiana “complex.” Van Haverbeke (1968a), through his study of these two taxa in the Missouri River Basin, has stated that: The entire ie in population within the Basin is apparently of hybrid derivation with neither of the extreme parental types being found. There is a trend of increasing hybrid index values (also d eod que plasm values) from ufu to northwest over the Basin from the reported range of J. virginiana to and into the reported range of J. scopulorum. This = may be the ita of bilateral introgression between the two species. There was, however, a strong tendency toward bimodality 8 the population as demonstrated by the res of two distributions in each of the three hybrid indices. This indicated the presence of two different species—J. scopulorum and J. virginiana. While Van Haverbeke (1968b) admits his data might be interpreted as 938 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 constituting evidence for introgression he, nevertheless, suggests, indeed cham- pions, an alternate hypothesis: As an alternative pU des) it would seem that because of the greater diversity of the jmipers in western North America, that J. virginiana was at some time derived from this area. It seems possible that with the inherent variability in the germ plasm ancestral to both J. POUR and J. virginiana, that propagules pas flourish in sites toward the east. This could have initiated an eastwar 5 propagu 5 through mutation and selection eventually became what we now recognize as J. vir, al inia — It should be noted that this latter evolutionary model is in direct conflict with that proposed by us (Flake, von Rudloff & Turner, 1969, 1973) in which we suggest that the Appalachian Region is the ancestral center for the origin of J. virginiana and its various races. Hence, the question of introgression between J. virginiana and J. scopulorum is left open by Van Haverbeke’s study. Initial investigation of the terpenes of Juniperus scopulorum, unlike that of J. ashei, showed that its volatile components were essentially those of J. virginiana, differing only in their quantitative expression. Subsequent populational analysis of the type employed in the J. ashei-J. virginiana studies showed that regional intergradation of the chemical characters occurred across the Missouri River Basin, much as found by Fassett (1944) and Van Haverbeke (1968a) for morpho- logical features. Three models might be proposed to account for the variation found in this region: l. ANCESTRAL GENE POOL—Juniperus scopulorum and J. virginiana may have arisen from ancestral populations largely endemic to the Missouri River Basin. Subsequent evolutionary divergence to the west and east, respectively, might have occurred, leaving a residuum of genes common to each in the area concerned. 2. ALLOPATRIC INTROGRESSION— The variability is due to extensive gene flow from J. scopulorum into J. virginiana as a result of hybridization and backcrossing in peripheral regions of contact and areas of sympatr 3. MIGRATORY TAILINGS—The River Basin was an ancestral migratory route through which J. scopulorum-like populations passed on their way to be- coming what is now known in the eastern United States as J. virginiana. In Van Haverbeke’s words (1968b), “Thus, rather than being considered as an intro- gressive series, this juniper population [those of the Missouri River Basin] can alternatively be interpreted as a divergent evolutionary series which has not yet completely separated." should be emphasized that in the investigation by Van Haverbeke about 40 morphological characters were selected for measurement and numerical analy- sis. These were obtained from some 700 trees from 72 sites scattered throughout the River Basin area. In spite of this excellently conceived, carefully documented, laborious study, the investigator was unable to decide, unequivocally, between models 2 and 3; in fact, he believed that his data best fit the migratory tailings model. (Model 1 was not tested, presumably because of its implausibility, con- sidering the biogeographic history of the Basin region.) Our own study (Flake, Urbatsch & Turner, 1978) also involved about 40 characters, all chemical. These were obtained from some 200 trees from 10 sites 1977] TURNER—CHEMOSYSTEMATICS AND TRADITIONALISTS 939 systematically selected at about 150-mile intervals in a southeast-northwest transect across the Basin. In spite of the fewer populations sampled and the smaller overall sample size, we conclude our data overwhelmingly suggest that the variable River Basin populations are the result of allopatric introgression, primarily in the direction of J. virginiana, much as Van Haverbeke thought might be the case, but the morphological characters which he used were not sufficiently indicative to prove decisive. MACROMOLECULAR APPROACHES If I were interested in obtaining the most meaningful arrangement of present-day angio- sperm families, phylogenetically speaking, I would rather have availa pis to me the primary structure ( amino acid sequence) of ten metabolically important enzymes (such as cytc home of all of the taxa which comprise these groups than have a detailed listing of all of the amarni features which characterize the groups (Turner, 1969) The d and proper taxonomic position of the hypothetical past organisms that repre- sent the branch points in the scheme cannot be determined solely from the dac iiid ane ‘of modern species as deciphered from the amino add sequences (Cronquist, 1976). Protein sequencing and other molecular methods ee a fact, become in the near future the most dua tools for the study of phlogeny ( Avala, 1976). The amino acid sequence trees are obviously more compatible with some possible phy- logenetic interpretations than others, or there would be no point in making them at all. we assumed that they were in all respects correct insofar as they go, they would place certain limits on the general phylogenetic trees that could be seriously dut, 1 1976 Though this be madness, yet there is method in't (Shakespeare, Hamlet, Scene II, Act 2). fossil evidence is highly in accord with an overwhelming mass of evidence from com- parative spen yor of living species that the Magnoliidae are the most primitive (i.e., st modified) group of ine angiosperms... (Cronquist, 1976). . the molecular tree indicates that present-day families represent relic groups which have for the most Pa rt had a long separate evolutionary history. They do not support the sug- gestion implicit in, for example, Cronquists scheme . . that the Magnoliidae gave rise to the Ç isanka idae on ihe one hand, and to the Rosidae on the other, the latter, in turn, giv- ing rise to the Asteridae ( Boulter, 1973) Something is rotten in the state of Denmark (Shakespeare, Hamlet, Scene IV, Act 1). With relatively few exceptions, the traditionalist might yawn at the seemingly trivial impact of micromolecular data upon his various systematic models. But he has not yet been able to treat with indifference the likely impact of macro- molecular data upon his most treasured erection, the "Tree" to plant families. As unrecognizable as this tree might be to the various workers concerned, any reinsertion of branches or elevation of roots, using such chemical data, is met with alarming cries from this or that proponent. I refer specifically to the recent paper by Cronquist (1976) entitled, "The Taxonomic Significance of the Struc- ture of Plant Proteins: A Classical Taxonomists View." This is a 27-page ram- bling review covering the whole field of comparative enzymology, the gist of which is, because these data do not or have not supported my particular views, there must be something wrong with the approach. The approach is the same as that which has been applied to animals success- fully, namely, the use of amino acids among the homologous proteins in different 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 organisms as an indicator of time of branching. And, strangely enough, he ac- cepts, in principle, the use of cytochrome c as a reasonable, but often unsteady, clock for animals, yet rejects this as valid for plants. I quote: Given the difference in evolutionary pattern between plants and a it should not be surprising if the animal iy rea aa system places stronger constraints on the acceptance of amino acid substitutions in cytochrome c than does the plant 8 system. It would be entirely in harmony with the other differenc ces in plant and animal evolution if the same kinds of changes could wi om by very different sorts of plants and if back mutations were not notably counter-select Cronquist focuses his attack largely upon the data from Boulter’s laboratory in Durham, England, which is the only group to sequence any significant num- ber of plant proteins, namely plastocyanin and cytochrome c. Amino acid se- quences from the latter, in particular, suggest that the familial tree is quite dif- ferent from the one proposed by Cronquist (and, of course, that of Takhtajan, the two being quite similar). This is disturbing: everyone should accept that the Magnoliidae among the angiosperms is primitive to everything else. He does not like the Caryophyllidae coming off as a first branch on the cytochrome c familial tree. He does not like to think of the morphologically highly advanced Com- positae represented as a very old isolated branch; everyone should know that it is recent, going back to the Miocene-Oligocene boundary (in spite of the fact that he acknowledges in footnote form that very recently published and unpub- lished pollen fossil data might push the family back to the Paleocene, if not ear- lier). To me, it is remarkable that a traditionalist of his stature, fully aware of the ubiquity of this macromolecule among organisms generally and cognizant of its crucial role in the metabolic pathway of both plants and animals, should at- tribute the discrepancies to poor or erratic functioning of this kind of clock, rather than to the morphological data, which, after all, has no face, no dial, no nothing to suggest the time of branching of this or that phyletic line. Cronquist (1976: 5), while accepting the general premise that the cytochrome c clock works for animals, nevertheless makes great gloat over the fact that the amino acid sequence of rattlesnake cytochrome c is out-of-line with the position of that organism in the phyletic tree. There follows a typical Cronquistian quote, “If the reported sequence for rattlesnake is correct, there seems to be no easy way to explain it, short of conjuring up a vision of a lonesome cowboy on the lone prairie, with none but a rattlesnake for company [referring to the seeming simi- larity of its sequence to that of the genus Homo]." I think that there are better ways to explain that single discrepancy, even if the sequence is correct. Cronquist presumably wants us to believe that an occasional unsteady tick (if even that!) in the animal world is sufficient reason to believe that this same clock is largely unsteady in the plant world. In his desire to discredit such data, at least that of cytochrome c, he likens this to the Age and Area concepts of Willis (ludi- crous! ), followed by the statement that: Evolution d other 5 in both plants and animals tends to undergo periods of rapid 5 interspersed with periods of more gradual change, and there is no a priori reason to suppose that "ern ‘significant changes in amino acid sequence would proceed any dif- 5 1977] TURNER—CHEMOSYSTEMATICS AND TRADITIONALISTS 24] Of course, that's the point; there has been a sufficiently long record of plant evolution so as to believe that the cytochrome c clock has some kind of accuracy; fast or slow upon occasion, it nonetheless seems to average out as stochastic over time. Anyway, a generally erratic clock is better than no clock, giving the muddle of morphological darkness within which most plant taxonomists work. As a final denouement, in case he hasn't convinced the reader, Cronquist adds a neat punch paragraph: This discussion of the evolutionary clock may be something like beating a dead horse, but some people are still trying to ride the horse. If the horse is really dead it won't mind the beatin One should perhaps remind Cronquist that, to judge from the recent articles by Fitch (1976), Zuckerkandl (1976), King (this symposium), and articles in press by yet other such workers, the horse is alive, is being ridden quite nicely, and perhaps doesnt deserve the beating being administered! It would be unfair to conclude this address with the audience feeling that Cronquist might be quite negative towards the application of chemical data to taxonomic problems. He is not, for he concludes, in hindsight, that: I welcome the appearance of amino acid sequences as an additional tool for taxonomists we have the sequences for several proteins from members of a wide range of fami- lies, weiding critically important ones, we can make good use of this powerful toc Lets hope he means this; in the meantime he might wish to s. Shakespeare’s King Richard the Third, “A dead horse, a dead horse, My Kingdom for a dead horse!” LITERATURE CITED Apams, R. P. B. Turner. 1970. L i pun and numerical studies of natural RR of ro s ashei Bush. Taxon 19: 728—751. ALSTON, R. E. & B. L. Turner. 1963. A on among four species of Baptisia Lor ituoxae V. Amer. J. Bot. 50: 159-1 ANDERSON, E. 1949. Introgressive Hybr XP John Wiley & Sons, New York. ————. 1953. Introgressive Hybridization. Biol. Rev. 28; 280-307. AYALA, F. J. 1976. Molecular genetics and evolution. Pp. 1-20, in P ]. Ayala (editor), Moleca Evolution. Sinauer Assoc. Inc., Sunderland, Massachuset Bouter, D. 1973. Amino acid sequences of pe sas c and Dlastdeya vanins in phylogenetic studies of higher plants. Syst. Zool. 9-553 CnowNQuisr, A. 1976. The Dp ais 818 Qa of the structure of plant proteins: a classi- cal taxonomist’s view. Brittonia 2 —27. Davis, P. H. & V. H. ck 1963. Principles of Angiosperm Taxonomy. Oliver and Fassett, N. C. 1944. Juniperus virginiana, J. horizontalis and J. scopulorum. Bull. Torrey , l - Frrcu, W. M. 1976. The molecular evolution of cytochrome c in eukaryotes. J. Molec. FLAkE, R. II. & B. IunxER. 1973. Volatile constituents, especially terpenes, and their utility and potai as taxonomic characters in populational studies. Nobel Symp. 25: 23-128. š UnnaArscH & B. TurNER. 1978. Chemical documentation of allopatric intro- gression in Juniperus oe and J. virginiana. (In pres ress, , E. vox RUDLOFF & B. L. TURNER. "ui 9. Quantitative study of clinical variation in Juniperus virginiana using terpenoid data. Proc. Natl. Acad. U.S.A. 64: 487—494. ———— & —. 1973. Confin e of a clin: j pattern of chemical differenti sion in Juniperus virginiana from terpenoid data ecd in successive years. Rece Adv. Phytochem. 6: 215-228, 242 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 LamisEev, M. M. 1940a. A monograph of the genus Baptisia. Ann. Missouri Bot Gard. 27: 119-258 1940b. Analysis of a hybrid complex between Baptisia leucantha and Baptisia viridis in Texas. Amer. J. Bot. 27: 624-628. Turner, B. L. 1969. Chemosystematics: recent developments. Taxon 18: 134-151. Van HavkEnBEKE, D. F. 1968a. A population analysis of Juniperus in the Missouri River Basin. Univ. Nebraska Stud., N. S., 38: 1-82. . 1968b A taxonomic analysis of Juniperus in the central and northern great plains. Proc. Sixth Centr, States Forest Tree Improv. Conf. 6: 48-52. WiLBun, R. L. 1963. The 1 plants of North Carolina. North Carolina Agric. Exp. Sta. Tech. Bull. 151: 1-294. ZUCKERKANDL, E. zvolutionary processes and evolutionary noise at the molecular l. II. A selectionist model for random fixations in proteins. J. Molec. Evol. 7: 269- SYSTEMATICS OF MORAEA (IRIDACEAE) IN TROPICAL AFRICA' PETER GOLDBLATT" ABSTRACT Moraea, with 24 species, is well represented in tropical Africa, although the center for e genus is to the south in the winter rainfall region of southern Africa, Nine species ar 1 for the first time. (M. callista Goldbl., M. afro-orientale Goldbl., M. iringensis Goldbl., M. inyangani Goldbl., M. angolensis Goldbl., M. tanzanica Goldbl., M. upembana Goldbl., M. a 7 M. balundana Goldbl. ) Only two of the five subgenera of Moraea occur in tre Africa, subgenus Vieusseuxia and subgenus Grandiflora, the latter in particular exh ibiting sinti radiation in south central Africa. On the basis of limited mowledge of 1 and cytological variation patterns in tropical Africa, Moraea is believed to be of recent origin here. Details of ecology, cytology, and evolution are elaborated and ceed to Pur cade in southern. Africa. Moraea is a large genus of some 95 species, occurring throughout sub-Saharan Africa. The genus comprises small to medium-sized herbaceous geophytes. It is found mainly in highland areas in the tropics, usually in well-watered grass- land, but also in open woodland or in marshy places. In temperate southern Af- rica, Moraea occurs at all elevations. Moraea is of some economic importance as all species are to some degree toxic to stock, section Polyanthes particularly so. Only a few species have been proven toxic, but all should be assumed to be until shown otherwise. The present revision treats only the tropical African members of the genus and will complete my taxonomic study of Moraea, begun in 1973 with a revision of Moraea in the summer rainfall region of southern. Af- rica, followed by a revision of Moraea in the winter rainfall region of southern Africa (Goldblatt, 1973, 1976b ) This treatment includes as tropical Africa the area south of the Sahara and north of the region covered by the Flora of Southern Africa [Namibia (South Vest Africa), Botswana, Lesotho, Swaziland, and South Africa]. In this area, extending from Rhodesia north to Nigeria and Ethiopia, there are 24 species of Moraea, a larger number than previously recognized, but low in comparison to the 76 species in southern Africa. Only five species occur in both tropical and the southern African summer rainfall region, while one species, M. spathulata, occurs in the winter rainfall region of southern Africa as well. The present work is the first modern comprehensive treatment of Moraea in tropical Africa and the only treatment since Baker’s (1898) revision for the Flora of Tropical Africa. RELATIONSHIPS The affinities and relationships of Moraea are discussed in detail in my re- vision of the species found in the winter rainfall region of southern Africa (Gold- This study was D ch By grant BM$-74-18905 from the U.S. National Scienc PINE I thank Jean Pawek for providing material for cytological study and for her ger erous assistance during field work in Malawi. I also thank Janet Klein for the illustrations iod my wi e Margaret for assistance in the field and with the manuscript * B. A. Krukoff Curator of African Botany, Missouri Botanical e 2345 Tower Grove Avenue, St. Louis, Missouri 631 10 ANN. Missouni Bor. GARD. 64: 243-295. 1977. 944 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 blatt, 1976b) and also in a paper dealing with the cytology and subgeneric classi- fication ( Goldblatt, 1976a). In the latter paper I expanded my earlier hypothesis that Moraea is a relative of the widespread northern hemisphere Iris and that these two genera probably had a common origin from ancestors of the genus Dietes. Moraea is, however, more closely related to a group of South African corm-bearing genera which, like Moraea, also have a secondary bifacial leaf than to Iris or Dietes. These genera include Homeria, Hexaglottis, Gynandriris, Galaxia, and Barnardiella which are placed with Moraea in Homerinae (Gold- blatt, 1976a), one of the three subtribes of Irideae, a predominantly Old World tribe of subfamily Iridoideae. GEOGRAPHY OF THE TROPICAL AFRICAN SPECIES Although Moraea occurs throughout sub-Saharan Africa, its present center is clearly the winter rainfall region of southern Africa which extends along the south and west coast of the Cape Province. Not only is this area richest with 54 species, 48 being endemic, but it is also the center of diversity in the genus. All five subgenera occur here and three of these are essentially endemic. The two largest subgenera Vieusseuxia and Grandiflora, also considered the more advanced, extend. well outside the winter rainfall region. In fact, the center for subgenus Grandiflora lies outside this area, in the mountains of southeast Africa, extending from Natal to Malawi, Tanzania, and Zaire. Subgenus Vieusseuxia, the second subgenus found in tropical Africa, com- prises two sections. Section Polyanthes, the less specialized, occurs from the southern Cape to Ethiopia and is well represented in Tanzania, as well as in the South African provinces of Natal and the Cape. The most primitive members of section Polyanthes are believed to be those with several leaves and unlimited branching, rather than single-leafed, few- or unbranched species; these include M. polyanthos and M. polystachya from the Cape Province and M. carsonii from Rhodesia-Zambia-Malawi. The center for this section thus may also lie in the mountains of southeast Africa but perhaps more likely to the south along the interface between the summer and winter rainfall regions. The second section of subgenus Vieusseuxia, section Vieusseuxia, is clearly specialized ( Goldblatt, 1976b) and does not occur outside southern Africa where it is concentrated in the winter rainfall region. In tropical Africa, Moraea occurs in almost all highland areas above 1,200 m and concentrations of species are found mainly in isolated areas that are con- siderably higher. Representation of the genus is notably poorer north of the equator with only two species in Ethiopia, three in the Uganda-Sudan-Kenya highlands, and only one in the mountains of eastern Nigeria and Cameroons. Significant areas of concentration are all in the highlands of central tropical Af- rica and several endemics occur in these isolated, and well-watered montane and semimontane regions. The main centers of endemism are as follows: 1. Inyanga highlands of Rhodesia: 5 species, one endemic (M. inyangani). 2. Northern Malawi-eastern Zambia escarpment and Southern Highlands of Tanzania: 8 species, 3 endemic (M. tanzanica, M. callista, M. iringensis ). 1977] GOLDBLATT— —MORAEA 945 3. North-central Zambia, southern and eastern Shaba: 14 species, 5 endemic (M. brevifolia, M. unifoliata, M. balundana, M. bovonei, M. upembana). Subgenus Vieusseuxia is predominantly eastern in distribution and though it occurs from Ethiopia to Rhodesia, it does not extend anywhere in tropical Africa west of longitude 24^E, and is thus absent from Angola and the Nigeria- Cameroon highlands. Almost as wide ranging as subgenus Grandiflora in tropi- cal Africa, iue m ö is poor in species, with only 7 species north of South Africa, and only 4 of these exclusive to tropical Africa. Significant speciation in nt ssp Vieusseuxia occurs only in the Southern Highlands of Tanzania where two very striking species, M. callista and M. iringensis, are en- demic. Other species of subgenus Vieusseuxia are widespread, with M. thom- sonii extending over almost the entire range of the subgenus in tropical Africa, and found from Ethiopia in the north to Transkei in southern Africa. Subgenus Grandiflora is well represented in tropical Africa by 16 species, only two also found in South Africa where an additional 11 species occur. The only really widespread species of this subgenus in tropical Africa is M. schimperi which extends over the whole range of Moraea in tropical Africa. Other species are considerably more restricted, and several species are known from single collections or a very small area. Moraea inyangani is one example while several species in the M. tanzanica-M. unifoliata alliance of reduced species are also very local, especially M. unifoliata, M. balundara, M. bovonei, all from Shaba, Zaire, M. brevifolia in northeastern Zambia, and M. tanzanica in southwestern Tanzania-northern Malawi. A very marked center of speciation for subgenus Grandiflora is evident in this belt across south-central tropical Africa. Hisronv The first collections of Moraea from tropical Africa were made as early as the 1840s when Schimper discovered M. schimperi in Ethiopia. Schimper's collections were first assigned to two separate species in the genus Hymeno- stigma. Knowledge was later extended by Welwitsch who collected M. schimperi. M. clavata, and M. textilis in Angola in the vears following 1853. Welwitsch's Angolan Iridaceae were only described much later, in 1878, by Baker who actu- ally admitted seven species from Angola including three now regarded as con- specific, and referred to Ferraria rather than Moraea. The important milestone in the knowledge of Moraea in tropical Africa is J. G. Baker’s (1898) treatment for Flora of Tropical Africa. Baker recognized 16 species in his treatment. However, four of these are clearly the same species of Ferraria, F. glutinosa, one is today regarded as Dietes, while of the remaining ll species M. welwitschii, M. zambeziaca, and M. diversifolia are conspecific ( M. schimperi) and another two, M. textilis and M. mechowii, are also regarded here as the same species. A last species, M. bella, was until recently associated with the Cape endemic M. angusta, although it is in fact unrelated. By 1898 only seven of the presently recognized species were known, six were named and one misidentified. Subsequent studies on Moraea were carried out independently by German, 246 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Belgian, and British botanists who confined their interests largely to the colonies of their respective countries. In particular, de Wildeman, working on the flora of what is now Zaire and Burundi, described six species, all new for the Congo, but four of which have proved to be synonyms of earlier species from other parts of tropical Africa. More recently, Moraea has received very little systematic at- tention in tropical Africa. In fact, the recent treatment by Geerinck (1970) for Zaire and Burundi is the only significant work in the last fifty years. Geerinck recognized six species, one tentatively; and subsequently one more (Geerinck, 972). Geerinck's species concepts were far broader than those held by me and I recognize several more species in the Zaire-Burundi area than Geerinck did. In the present revision 9 of the 24 species are new to science. Knowledge of the genus in tropical Africa is, however, far from satisfactory, and several species are known from one locality while collections of some species are incomplete, lack- ing corms or fruits or adequately preserved flowers. ECOLOGY HABITAT Moraea occurs in two distinct types of habitat in tropical Africa, either in dry situations in grassland and open woodland or in wet, marsh situations known in southern Africa as vleis, and in south tropical Africa as dambos. Dambos may be seasonal or permanent, but have the characteristic of being poorly drained yet seldom deep, and usually have a rich vegetation including many grasses, sedges, as well as geophytes and herbaceous dicots. The dry habitat may com- prise more than a single niche for Moraea, but data available does not indicate this, rather suggesting that species may be fairly tolerant of minor ecological differences and thus growing equally well in open woodland or grassland of various types. Most species, though not all, appear to be restricted to either one or other of the two major habitats. Records suggest that M. ventricosa and M. textilis are exceptions, as they are listed as having been collected in wood- land, grassland, or on the edges of marshes. Personal observation has shown that M. schimperi also occurs from wet marshes to open, well-drained grassland, the latter habitat being occupied only in areas of higher rainfall. FLOWERING TIME Species of Moraea may be found in bloom almost throughout the year, though few species flower in the dry season. Peak blooming time is mid wet season. Each species, however, has its own fairly limited flowering period, and detailed examination of flowering times suggests that this factor is of considerable sig- nificance in the evolution of the genus in tropical Africa. In general, it appears that only a few species, and only one in each subgenus, may bloom in a particu- lar habitat in any given locality at one time. Species of Moraea tend to bloom for about two months; thus in a particular area several species might occur, each flowering at a different time. The significance of flowering times is elaborated further in the following discussion on evolution. 1977 GOLDBLATT—-MORAEA 947 EVOLUTION The pattern of species that prevails in Moraea in tropical Africa suggests that the group is rapidly evolving into a limited number of spatial, temporal, and ecological niches available to the genus. A very limited degree of important floral differences suggests that there has not been sufficient time for fundamental morphological changes to become established in the flowers, so that selection for different pollinators has not been accomplished. There are three major groups which are dealt with separately in the follow- ing discussion: the large-flowered M. spathulata-M. textilis series of subgenus Grandiflora; the smaller flowered and morphologically specialized M. tanzanica- M. unifoliata series of subgenus Grandiflora; and the small uniformly blue- flowered subgenus Vieusseuxia. The Moraea spathulata-M. textilis series. The series comprises vegetatively similar, large-flowered species in which both flowering time and habitat differences have played the major role in their evolution. The dry habitat does not support species of subgenus Grandiflora in the dry season, with the possible exception of the poorly known September-blooming M. upembana. Early in the wet season, from December, M. verdickii blooms in Tanzania, Zambia, and Zaire. Moraea ventricosa follows in this region with a blooming peak in March. Subsequently, at least in the east, in Malawi and Tanzania, M. macrantha blooms from March to June, lasting into the beginning of the dry season. In the wet habitat, the wide-ranging M. schimperi blooms from September to early December south of the equator. It is replaced later in the season by a 'ariety of more localized species of the series dealt with in the following section, including M. brevifolia in Zambia, M. unifoliata, M. bovonei, and M. balundana in Zaire. Later, towards the end of the wet season, M. bella blooms in Zambia, Zaire, Malawi, and Tanzania. Geographical isolation is of relatively minor significance in the evolution of this series but has played a role as in M. ventricosa-M. textilis, which occupy similar habitat and temporal niches but have different geographical ranges. A similar role for geographical isolation is seen for M. verdickii-M. spathulata. 2. The Moraea tanzanica-M. unifoliata series. The species of this series are all relatively small in size, have similar vellow flowers, and exhibit distinctive vegetative modifications making them easy to recognize. Unlike the previous series, all, with the exception of the poorly un- derstood M. upembana, occupy a similar habitat and flower at the same time. Spatial isolation is the major factor of evolutionary significance here, with each species relatively limited in range and none sympatric. Small, local populations are characteristic in the series. 3. Subgenus Vieusseuxia. Subgenus Vieusseuxia, also a group where species have very similar flowers. exhibits a similar pattern of evolution to subgenus Grandiflora with ecological factors assuming dominance over a secondary geographic element. The dry 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 — grassland habitat supports M. thomsonii in the later dry season; Af. carsonii in the early to mid wet season at least in the south; M. afro-orientale in similar situ- ations north in Tanzania, Kenya and Uganda; and M. iringensis locally. In the wet habitat, M. natalensis blooms in the wet season throughout south tropical Africa. There are no late-blooming species and presumably taller grasses shade out these smaller plants in late season. The exception to the patterns described above is the very striking M. callista. This species has à large and very different flower from its allies and, although it presumably evolved in isolation, it is now distinguished by its floral peculiar- ities and, by inference, has a distinctive series of pollinators. Summarizing, two main modes of evolution appear to have been operative in the tropical African species of Moraea. These are: (1) an ecological factor (habitat and flowering time) and (2) a geographical (spatial) factor. Within the two major groups, subgenera Grandiflora and Vieusseuxia (excluding M. cal- lista), floral differences are small and probably not of adaptive significance, so that the species of each group do not appear to have evolved in response to pres- sures from pollinators. This situation is similar in the summer rainfall area of southern Africa, al- though somewhat more complex, with more species and basic types, and an ele- ment of floral evolution is evident in section Vieusseuxia. However, the evolu- tionary pattern in the winter rainfall region is quite different. Flowers vary considerably and floral adaptation is of the greatest significance. Evolution for limited and specific ranges of pollinators consequently must have been im- portant. Flowering time and habitat are minor as most species grow in essentially similar situations and all are spring blooming. Soil differences assume great importance as does the geographical factor. Related species are often isolated from one another geographically or by distinct soil preferences. The evolutionary patterns here, in fact, are typical of an area with a Mediterranean climate (Raven, 1973) TAXONOMIC CHARACTERS The morphology and important taxonomic characters were dealt with at length in my recent paper on Moraea in the winter rainfall region of southern Africa and little need be added here. It will, however, be helpful if I summarize the important features of the two subgenera that occur in tropical Africa. Subgenus Grandiflora comprises the largest-flowered species in the genus, though a few specialized species in tropical Africa—e.g. M. clavata, M. angolen- sis, and M. unifoliata have relatively small flowers. Floral morphology is uni- form apart from size and color, most species having yellow flowers, only M. schimperi, M. ventricosa, M. macrantha and M. textilis having blue flowers. Outer tepals are outspread, the inner = erect. All species are unbranched, and have a single leaf, usually well developed and basal, but in a few species re- duced and inserted well above ground level. All species in which fruits are known have large flattened + discoid seeds and large capsules. The subgenus extends from the southern Cape in South Africa to Nigeria and Ethiopia but 1977] GOLDBLATT-MORAE A 249 Chromosome number in tropical African Moraea. An asterisk (*) indicates a count for South African material only; new counts are in bold type Diploid number 2 Species 9n zollection data Or reference M. carsonii 12 Goldbl: att, 1976a V. elliotii 12* 24* Lewis, 1966; Goldblatt, 1976a. M. natalensis (as M. erici-rosenii ) 12 Lewis, 1966. M. thomsonii (as M. stricta) 12 Chimphamba, 1974. 48 ee Zomba Mt., Goldblatt s.n. ) voucher 94*, 36% Colaba 1971 M. spathulata (incl. subspp.) 12* , Goldblatt, 1971, 1976a. M. schimperi 12 Goldblatt, 1971; are he 1974. Malawi, Zomba Goldblatt 4259 (MO ). Malawi, ^ ae Mzu- Goldblatt 4590 (MO). Malawi, 53 MER Marymount, Patek 583: ) M. macrantha (as M. textilis) 12 Goldblatt, 1976a. 12 Malawi, Mzuzu, Pawek 5396 ( MO). Malawi, | Katumbi-Nyika — intersec- tion, Goldbl att. s.n. (no voucher ). M. ? ventricosa (flowers not seen) 12 Burundi, Mt. Bona, E. Bujumbura, Goldblatt s.n. (no voucher). M. tanzanica 12 Malawi, Nyika Plateau, unies for- est road, Pawek 6660 (N )). is notably absent from the southwestern Cape, an important center for the genus as a whole. Subgenus Vieusseuxia, composed of two sections of which only section Poly- anthes occurs outside South Africa, comprises small to fairly large plants all with very similar blue (to white) flowers, which in tropical Africa are quite small and in marked contrast to subgenus Grandiflora. Tepals of both whorls are out- spread, the exception being M. callista from southwestern Tanzania which has unusual, large blue and white flowers with reflexed tepals. Species varv in leaf number and position of insertion which ranges from basal to just below the in- florescence in M. natalensis. Moraea carsonii has 2-3 leaves, M. callistra has two, while other species typically have one leaf. Seeds of all species, where known, are small and angular and capsules are. small. The subgenus extends from the southwestern Cape to Ethiopia but is absent from West Africa and An- gola. Section Polyanthes is the more widespread, with section Vieusseuxia re- stricted to Africa south of the Limpopo. CYTOLOGY Moraea is well known chromosomally (Goldblatt, 1976a), some 65% of the 95 species having been studied. It is least well known in tropical Africa and only 9 of the 24 species have now been counted. Counts for the genus in tropical Af- rica are summarized in Table 1, where several new counts are presented. 950 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 The tropical African species have fairly uniform karyotypes, with a base number of x — 6. The chromosome complement in all species of subgenus Grandiflora examined is similar and all species are diploid, 2n — 12, with 4 large telocentric pairs, and two = acrocentric pairs. Satellites are located on a telo- centric pair ( Goldblatt, 1976a: 13). Moraea schimperi is distinctive in having a secondary constriction on the end of the long arm of an acrocentric pair, a feature also noted by Chimphamba (1974) n subgenus Vieusseuxia the chromosomes are acrocentric with the longest pair almost metacentric (Goldblatt, 1976a: 13). Polyploidy has been reported in both sections. In section Polyanthes, M. thomsonii is heteroploid with tetraploid and hexaploid plants recorded in South Africa and an octoploid from Malawi. Chimphamba (1974) has reported 2n — 12 in this species but determination needs to be confirmed and the locality is unknown. Moraea elliotii is also hetero- ploid, with diploidy and tetraploidy reported in South African plants. The cytological uniformity of Moraea in tropical Africa, especially in sub- genus Grandiflora, contrasts with a great degree of heterogeneity in South Af- rican winter rainfall area species. This uniformity supports my suggestion based on limited morphological variation that Moraea is of fairly recent origin in tropi- cal Africa. TAXONOMIC TREATMENT? Moraea Miller, Figs. Pl. 159, tab. 238. 1758, as Morea and altered to Moraea by Linnaeus, nom. cons. TYPE: M. vegeta L. For complete synonymy and generic description see Goldblatt (1976b). Distribution: Sub-Saharan Africa, in highland areas in the tropics, at all al- titudes in temperate southern Africa; species concentrated in South Africa, mainly in the southwestern winter rainfall region. KEY TO THE SPECIES 1. Produced leaves 2 or more, well developed. 2. Sheathing portion of upper (cauline) leaf more than 1.5 cm long. 3. Stem few to many branched; spathes = cm long ue ee a 1. M. carsonii : Stem simple or 1-branched; spathes ride n long 2. M. callista athing portion of upper leaf 2-6 mm I. afro- “orientale i? Produced leaf solitary, occasionally reduced a +bractlike or pus i flower im 4. Plants either leafless | s EUIS time, or with a dry withered leaf attached; os in the dry se : wers yellow; isi, lens ils 2.7-4.5 cm lon 21. M. upembana Ç 15 pale blue violet; outer tepals 1.5- 2.2 cm long 7. M. thomsonii * All major collections of African flora were consulted for this study, and unless other- wise stated all type = cimens were seen. Field work was carried out in Malawi and to a iir bs e in Rhodesia. cimens examine ved are generally arranged in accordance with modern floristic projects -= particular areas. Modern place names are used, but old names where well known ently altered, are alt in parentheses. Collection data or liter: ature citation for chromosome numbers in the descriptions are given in the section on CYTOLOGY 1977] GOLDBLATT— —MORAEA 95] ' Plants with a green produced leaf at flowering time, — well developed but eee very short and inserted just below the in florescene eaf inserted immediately oS the inflorescence (leaf sometimes bract- ike and almost entirely sheathing ). Flower blue; stem arali branched 5. 6. M. natalensis 7. ' Flowers vellow; stem simple. Outer inflorescence spathe only slightly shorter than the inner. Leaf barely distinguishable from the spathes and not or onl shortly exceeding them 2.241. M. unifoliata 9.“ Leaf several times longer than the spathes 2... — 22. M. bovonei Outer inflorescence spathe less than half as dem as the m. OSEE A A AE: balundana 6.’ Leaf inserted from the base to the upper part ‘of the stem but ade rione ately below the inflorescence 10. Outer tepals 1.4—3.5 cm long. Sheathing portion of bract leaves at least 1.5 cm long. Flowers blue; outer tepals 1.4-2.5(—3.0) cm long. 3. Leaf inserted in the upper third of the stem 6. M. natalensis 13.’ Leaf inserted in the lower third of the stem 5. M. elliotii 12. ae » yellow; outer ak (2.0—)2.5-3.5 cm rns. : ict. leaf solitary 9 14. Bract leaves 3 or 15. Spathes 5-8 cm long; bract leaves 5-8 cm long — — BUM. . M. inyangani 15.’ Spathes 4.5-5.5 em Jong; bract leaves 3.0-4.5 em 1 » t — i . clavata ng ME T . 19. M. angolensis 11.’ Sheathing portion of bract leaves 1-7 mm | long. 16 3-5 mm long; anthers 4-5 mm long — 3. M. afro-orientale 6. ' Ovary 7-13 mm long; anthers +6 mm long — 4. M. iringensis 10.“ Outer tepals 3.8-10 cm long. 17. Bract and spathes ey at flowering time and the flowers blue and blooming at the a 2 = dry season to early in the wet sea- son (Sep.-Dec. sout equator, Dec.-May north of the equator); leaf often 1 than the stem, the e dark »own and prominen M. schimperi Bracts and spathes at least partly herbaceous at 1 time, or if not, the flowers white to yellow and not flowering at the end of the dry season; leaf exceeding the stem, or if not, flowers white to yellow, 18. Leaf inserted above ground level, very short and rarely ex- ceeding the "pee, the bract leaves usually oat : M. brevifolia 18.’ Leaf pem d Or if buena: well. abovg dme pue] then the za bract leaves 1 or 2, the j aves not exceeding the spathes or the pants no more than 35 cm high. 19. Plants of Rhodesia, southern Mozambique (and South rica 20. Flowering in spring and early summer, Sep.—Nov usually in moist habitats; 30-50(-70) cm tall; eaf ca, 5 mm wide. 10. M. muddii 20. Flowering in summer E open essay: unis more than 50 cm tall; leaf usually more than 1 c wide 8. M. mitis 19.' Plants of Angola and occurring ini of Rhodesia and southern Mozambique. 2]. Plants 30-35 cm tall with 2 bract leaves only and the bat only shortly or not exceeding the spathe . e 18. AT. tanzanica 91,’ Plants nalis more G han 40 cm tall, if less, then the bract leaves more than 2 and the leaf nud exceed- ing the spathes. 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 22. Bracts not overlapping. 23. Flowers blue. 24. Outer tepals 5.5-8 cm long; corm tu- nics of pale, reticulate fibers; ng of Zaire, Tanzania, Zambia, Malaw 5. M. macrantha 24.' Outer tepals 4.7-6 c n long; corm tu- nics of wiry, dark fibers; aes of An- 17. M. textilis go 23.’ Flowers. yellow Flowering Neo ov. to Feb. (Mar.), usu- ally in open grassland; anthers 10-14 mm long; outer tepals 5-10 cm ` e. UU 7 7 7 7 7 T 14. M. verdickii 25. Flowering (Feb.) Mar. to July in damp situations; anthers 8-10 mm long; outer tepals 4.5—5.5(—6.5) cm 13. M. bella Br acts ov erlapping Flowers blue. 27. ete tepals more than 5.5 cm long 15. a macrantha 27.’ Outer 97 85 less than 5.5 cm lo ong. 28 nthers 8-10(—11) mm » long in- ner cal 3.7-4.5 long; pa = Zaire, Zambia, 'Burur adi, M. ventric osa 28. wes (1010. 5-15 mm long; inner tepals 4.8-6.5 cm long; plants of Angola 17. M. textilis 26.’ Flowers white to yellow 29. Anthers 8-10 mm long; bract number 16. M. ventricosa 3-4(-5) . 29.“ Anthers 10-15 mm m long. 30. Outer tepals 5-10 cm long; bract leaves 2-3(or 4 jx then the tepals more than 5 cm) _ mn es 14 M. verdickii 30. Outer “tepals 4.5-6.5 cm long; bract leaves (4—)5-7 17. M. textilis Subgenus VIEUSSEUXIAA (de la Roche) Baker—Section Potyantues Goldbl. l. Moraea carsonii Baker, Bull. Misc. Inform. 1894: 391. 1894. type: Zambia, Mbala (Abercorn) district, “Fwambo,” Carson s.n. (K, holotype ).—Fic. IA. M. silanes De Wild., Contr. Fl. Katanga, Suppl. 4: 7. 1932. TYPE: Zaire, Shaba (Katanga), r Kolwezi, Homblé 1024 BR, holotype). — Plants small to medium in size, 20-40 em high, usually bearing several branches. Corm ca. 1.5 em in diameter; tunics of dark brown, fine to medium reticulate fibers. Prophylls dry and papery, pale brown, the upper often torn somewhat distally, occasionally becoming fibrous. Leaves 2, occasionally 3, the lower basal or inserted well below the branches, the upper cauline, canalic- ulate, 2-5 mm wide, usually falcate, exceeding the infloresence; sheath of upper leaf 1.5-2 cm long. Spathes occasionally reddish, herbaceous below, be- coming dry from the apex, the margins dry, pale brown, the apices dark brown, attenuate; inner spathe 3.2-5.0 cm long, the outer about two-thirds the inner. FIGURE 1. GOLDBLATT—-MORAEA Moraea species.—-A. M. carsonii, —B. M. afro-orientale. ( x0.5). 954 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Flower blue with yellow nectar guides; outer tepals 2.1-3 cm long, lanceolate spreading, the limb slightly longer than the claw; inner tepals 2-2.8 cm long, spreading. Filaments ca. 7.5 mm long, free in the upper third; anthers 4.5-5.5 mm long. Ovary 3.5-7 mm long; style branches 8 mm long, the crests 7-10 mm long. Capsule ovoid, to 8 mm long; seeds small, angular. Chromosome number 2n — 12 Flowering time: December to February. Distribution: Rhodesia, Zambia, Malawi, southern Zaire.—Fic. 2. Its two or sometimes three leaves, indeterminate growth pattern with several to many branches and small unspecialized flowers, place M. carsonii in a primi- tive position among the tropical African species of the genus. It clearly belongs in subgenus Vieusseuxia section Polyanthes and is probably most closely allied to the South African, southern Cape species, M. polyanthos, which generally has three leaves and similar flowers, and also to the widespread southern African M. polystachya, a taller species with much larger flowers. Moraea carsonii ap- pears to represent a link between these two multi-leafed southern African species and the several tropical and South African summer rainfall region species of section Polyanthes, most of which have a very similar flower but only a singie leaf, and are distinguished mainly by differences in growth form. Moraea carsonii has a relatively wide range, occurring in the Zaire province of Shaba (Katanga), in northern and eastern Zambia, and adjacent Malawi. A somewhat different form is found to the south in the Inyanga highlands of Rho- desia, which is generally more robust, and has larger flowers, giving it a resem- blance to M. polystachya to which it is frequently referred. Not all plants from Inyanga are equally robust, and several collections cannot be distinguished from plants occurring further north. For this reason the Inyanga populations are not accorded taxonomic recognition here. In Tanzania and northwards to the Sudan M. carsonii is replaced in EI habitats by M. afro-orientale Goldbl., which has until now been included in carsonii (Burtt, 1938). Moraea afro-orientale differs in having a single xs leaf inserted below ground, smaller flowers, and fewer branches which are often clustered in umbellate fashion. Occasionally, particularly in southern Tanzania, forms occur with two leaves but the upper leaf has an unusually short sheath. MALAWI. NORTHERN REGION: Mtwalo-Mzimba road, Pawek 3358 (K, MAL). Vipya hills S of Mzuzu, Pawek 4421 (K). SW Mzuzu, Pawek 8051 (MO, SRGH). 5 km S Mt. Hora, "mpl district, Hillard & Burtt 4456 (K, MAL, SRGH). CENTRAL REGION: Kongwe Mt. ne „ Robson 1654 LEM, LISC, PRE, SRGH Rn IODESIA. EASTERN REGION: Mt. Inyangani, Plowes 2158 (PRE, SRGH); Davidse, Simon & Pope 6536 (MO); Norlindh S Weimarck 4968 (BM, BR, K, PRE, SAM); West 7010 (S RCH); Wild 4927 (K, LISC, MO, PRE, SRGH). Near Bor nda Mission, . 8 55 5478 COI, K, MO, PRE, SRGH Above Rhodes Hotel, Inyanga, Whellan & 997 (SRGH). Nyamaropa Forest Reserve, Inyanga, Dale SKF204 (K, SRGH); 2 8 28, 54 (SRGH); Wild 7495 (BR, K, LISC, PRE, SRGH, MO). Inyanga Downs, Wild 5475 (K, MO). — Zai HABA: 10 km S of Lubumbashi (Elizabethville), Schmitz 1286 (BR). Lubum- bashi, Sale ^siens 1062 (BR). Fu ingurume, Symoens 14023 (BR, K). ZAMBIA: CENTRAL REGION: Lusaka, Angus 1466 (K, LISC, SRGH). cae River, Allen 498 ( K, SRGH). Kundalila Falls, Serenje district, Strid 2900 (K). NORTHERN REGION: Mbala township, Sanane 986 (K). Near Nakatali, Richards 8023 (K). abe "Falls, Mbala 1977] GOLDBLATT- -MORAEA 955 — (Abercorn), 7 8 3933 (K). Sansia Falls, a district, Richards 7436 ae Mbesuma ranch, Astle 5 (K, SRGH). WESTERN REGION: Luanshya, Fanshawe 1739 (BR, K, SRGH ). Mufulira, M 11749 (SRGH). Ndola, Pan Bange 612 (BR, K) 2. Moraea callista Goldbl., sp. nov. tyre: Tanzania, Southern Highlands, El- ton Plateau, Richards 7562 (K, holotype; BR, isotype ).—Fic. Planta 30-70 cm alta. Cormus ignotus. Folia duo, canaliculata, inferius 1 caulis simplex vel uniramosus. Spatha inflexae, herbaceae, exterior 4-6 cm longa, interior 34.5 cm longa. Flores caeruleo-malvini, tepala albe 'scentes distale; tepala exteriora 3-3.5 c ted imbis 2-2.5 em longis, reflexis; interiora breviora, reflexa. Filamenta ad 8 mm longa; EE 6 mm longae. Germen 5-7 mm longum; rami styli ca. 8 mm longi; cristae 5-7 mm longae. Plants solitary, simple or rarely 1-branched, 30-70 cm high. Corm not known. Prophylls membranous, the uppermost dry and fibrous towards the apex. Leaves 2, the lower basal and larger, the upper cauline, to 7 mm wide, canaliculate with slightly thickened hyaline margins, as long as, or slightly exceeding the in- florescence. Spathes often inflexed, herbaceous, with dry upper margins; outer spathe 4-6 cm long, the inner ca. ? the inner. Flowers blue mauve with the tepals fading to white distally; outer tepals 3-3.5 cm long, the limb 2-2.5 cm, fully reflexed when open; inner tepals somewhat smaller, also reflexed. Fila- ments to 8 mm long, united in the lower 5 mm; anthers 6 mm long. Ovary 5-7 mm long; style branches ca. 8 mm long, very broad and outspread, the crests 5-7 mm. Capsule and seeds unknown. Chromosome number not known. Flowering time: January to February in the west, May in the east. Distribution: Southwestern and eastern Tanzania in mountain grassland. 1,800-3,000 m.—F ic. 2. G Moraea callista stands apart from the other species in section Polyanthes be- cause its large striking flower with blue and white, fully reflexed tepals are quite distinct from the usual small, uniformly blue flowers with spreading tepals of other species. It occurs in two widely separated areas, high mountain grass- land in the Njombe area in the southwestern part of Tanzania, and to the east in the Uluguru mountains where only one gathering has been made, flowering at a much later date, in May, compared to January and February in the western part of its range. The species is known from very few collections and is all too little understood. TANZANIA. SOUTHERN HIGHLANDS: Elton Plateau, Procter 1646 (EAH); Richards 7562 (BR, K). Mangale-Njombe road, Richards 14216 (K). Mwake 12 ogg district, Richards 7823 (K). MOROGORO DISTRICT: Mzumbi, Semsei 1707 (EAH, K, PRE). 3. Moraea afro-orientale Goldbl., sp. nov. Type: Uganda, Northern Province, Mt. Debasien, Hedberg 1953 (UPS, holotype; EAH, K, S, isotypes).—Fic. IB. Planta 15-40 cm alta, gracilis. Tunicae cormi brunneae, reticulatae, tenuis. Folium solitarii, basale, raro folio secundo caulino, canaliculatum. Caulis ramosus, ferens um 0 olio caulinum unum, vagina brevissima. Spatha . ny rior 2-3.5 cm longa, exterior 1-2 mm brevior. Flores caerulei; tepala 2 .8-2.6 cm longa, d tepala interiora 1.5-1.8 cm longa. Filamenta ca. 5 mm longa; k e £ ca. 4. 5 mm longae. Germen ad 8 mm longum; rami styli ca. 5 mm longi; cristae ad 5 mm longas. Plants small, solitary, usually few branched, 15-40 cm high. Corm ca. 1.5 cm in diameter; tunics brown, cancellate to reticulate, of medium to fine fibers. 956 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 [ 20 40 f J — 9 30, LÀ Fg 4 20 |. L d 1 l 8 — at ^ f yf í ; SY IN Si / ( — Y s 3 10 A c2) l al inte Reha: ) tS po \ ba | V ! 1 V. 0 0 10 | i l @ W. carsonii Å v. callista 20 @ M. afro-orientale | A u. iringensis \ 80 0 100 200 300 400 $00 000 wiLES TS — n p—ssnJ Ho T — ns J O 200 400 800 800 1000 KM | i . — — + { | | | | 20 10 l 0 10 20 30 "m 50 ) | — 1005) Ficure 2. Distribution of Moraea carsonii, M. callista, M. afro-orientale, and M. irin- gensis. Prophylls usually 3, membranous, entire, obtuse-truncate. Leaf usually soli- tary, basal, inserted below the ground, the base covered by prophylls, canalic- ulate, 2-5 mm wide, relatively short, as long as or slightly exceeding the in- florescence; occasionally a second leaf developed at the first aerial node, usually 1977] GOLDBLATT—-MORAEA 957 bractlike, but 2-10 cm long, occasionally also exceeding the inflorescence; cauline leaf sheath 3-6 mm long. Spathes subequal, herbaceous, the margins mem- branous, rarely dry but not brown, the apices acute, brown tipped; inner spathe 2-3.5 cm long, the outer 1-2 mm shorter. Flowers blue with orange nectar guides; outer tepals 1.8-2.6 cm long, lanceolate, the limb slightly longer than the claw, spreading; inner tepals 1.5-1.8 cm long. Filaments ca. 5 mm long, free in the upper third; anthers ca. 4.5 mm long. Ovary 3-5 mm long; style branches ca. 5 mm long, the crests to 5 mm. Capsule ovoid, to 8 mm long; seeds small, angular. Chromosome number not known. Flowering time: April to May north of the equator in Kenya, Uganda, and Sudan; December to January south of the equator. Distribution: Southern Sudan, western Kenya, Uganda, and central and western Tanzania; in short grass and often in seasonally waterlogged ground, blooming soon after the start of the rainy season.—Fic. 2. Moraea afro-orientale is closely allied to M. carsonii, from which it is prob- ably derived, and it has until now been included in this species. It differs, how- ever, in several respects and is distinctive in overall habit, with prominent mem- branous prophylls, short stem and few, often clustered branches. Most forms have only a single comparatively short and erect leaf, inserted below ground level in contrast with the two long, flaccid leaves in M. carsonii. In the southern part of its range, in central and western Tanzania, M. afro-orientale often has two leaves, but when this is the case, the upper, cauline leaf is always distinctive in having a very short sheath (2-5 mm), contrasting with the more usual longer sheath (10-20 mm) in M. carsonii and its other relatives. In the single-leafed northern forms of M. afro-orientale the second leaf is reduced to a bractlike structure, often with a short free apex, and this bract also has a very short sheath. Moraea afro-orientale shares this distinctive, very short leaf or bract sheath with two other tropical African species, M. iringensis, which has a larger flower and a long ovary which is included in the spathes, and M. callista, which has two large leaves, and a large striking flower with blue and white, fully reflexed tepals. Both these species occur rather locally in southwestern and central Tan- zania on the southern extremity of the range of M. afro-orientale (Fig. 2). NYA. NYANZA PROVINCE: Bungoma district, Kokwaro 134 (EAH). SE slopes of Mt. Elgon, Padwa 16 (EAH, F, K, PRE, SRGH). RIFT vALLEY PROVINCE: Elgon and Trans Nzoia, Tweedie 444 (K). Kitale- Suam Mill road, Bally 2487 (K). Slopes 1 5 5 Endebe Polhill 411 (BR, K, PRE). * 3 545 (K); Lugard 573 (K); Adamson 513 H, K). Endebess, ct Pta ie a H). SUDAN. EQUATORIA: Mt. Lotuke, ium Mts., Jackson 1333 (BM); MacDonald 91 (BM); j Ayers 10952 (K IZANIA. CENTRAL PROVINCE: Singida-Dodoma, Feirarzl 5708 (EAH). SOUTHERN HIGHLANDS: Igurussi, Mbeya district, Procter SS (EAH, K). Ruaha National Park, Ma- gangwe Hill, Bjornstad 2231 (EAH, K). WESTERN PROVINCE: Moanzi, Sumbawanga dis- trict, Vezey- oe 1378 (K, SRGH). ES Lukunga confluence, Ufipa district, Rich- ards 10259 (BR, K). Above Malonje, near Sumbawanga, Richards 7 7210 (K BUGANDA PROVINCE: Mengo, Dradu 846 (EAH). EASTERN PROVINCE: Butiro, Licbenberz $46 (K). NORTHERN PROV ne E: Karamoja district, Lodoketeminit, near Moroto, Kerfoot 995,4947 (EAH, K). Mt. Debasien (Kadam), Hedberg 1953. (EAH, K S UES Moruita, Eggeling 5785 (BR, K). Karamoja, Thomas 2836 (BR, K). = 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ~ A. M. callista.—B. M. iringensis.—C. M. natalensis. Ficung 3. Moraea species. (0.5 4. Moraea iringensis Goldbl., sp. nov. Tyre: Tanzania, Southern Highlands, Kyimbila-Tandala, Stolz 2362 (K, holotype; BM, BR, PRE, isotypes. )—F ic. 3B Planta 15-30 cm alta. Folium solitarium, basale, usitate parum inflorescentium brevior, canaliculatum. hele pa pauciramosa ferens bractea una vaginis brevissimus. Spathae sve ts interior 3—4.5 cm longa, exterior 1.5-2.3 cm longa. Flores caerulei; tepala exteriora ca. 3 cm longa, effusa; ea ca. 2 em longa, effusa. Filamenta ca. 7 mm longa; antherae ad 6 mm 1977] GOLDBLATT—-MORAEA 959 longas. Germen 8-12 mm longum, non exsertum; rami styli ca. 1 cm longi; cristae ad 8 mm longas. Plants solitary, (10-)15-30 cm high, usually branched. Corm ca. 1 cm in diameter; tunics of fine to medium reticulate fibers. Prophylls 2-3, entire, mem- branous, often red-flushed, obtuse-truncate. Leaf solitary, basal, usually slightly shorter than the inflorescence, canaliculate, to 6 mm wide, with slightly thickened hyaline margins. Bract leaves to 3 cm long, sheathing at the base only, acuminate. Spathes herbaceous, acuminate; inner spathe 3-4.5 cm long, the outer + half the inner, often with the upper part not sheathing. Flower blue; outer tepals ca. 3 cm long, lanceolate, the limb ca. 1.5 cm long, spreading to slightly reflexed; inner tepals ca. 2 cm long, spreading. Filaments ca. 7 mm long, united in the lower third; anthers 6 mm long. Ovary unusually long, 8-12 mm, enclosed in spathes; style branches ca. 1 cm long, the crests to 8 mm long. Capsule and seeds not known. Chromosome number not known. Flowering time: December to January. Distribution: “Grassland and open Brachstegia woodland” in the southern highlands of Tanzania, recorded between Sao Hill and Makumbako, 1,800-2,200 m.—Fic. 2. Moraea iringensis is closely allied to the more widespread M. afro-orientale which occurs in western and central Tanzania, Uganda, western Kenya and the southern Sudan. It differs in being more robust, with larger leaves and flowers, and it is distinctive in having a very long ovary, 8-12 mm long, compared to 3-4 mm in M. afro-orientale. In spite of its length, the ovary is almost entirely enclosed in the spathes, in contrast to the exserted ovary found in all allied species. 'ANZANIA. SOUTHERN HIGHLANDS: Sao Hill, Iringa district, Chambers 38 (EAH, K); Carmichael 331 (EAH); Robertson 825 (EAH). Kyimbila, Tandala, Stolz 2362 (K, BM, BR. RE). Mufindi, Bjornstad 577 (K). 20 km N of Sunji, Njombe district, Sturtz 79 ( DAR, 9. Moraea elliotii Baker, Handb. Irid. 58. 1892. type: South Africa, Transvaal, marshes near Lake Chrissie, Scott-Elliot 1592 (K, holotype). For complete synonymy see Goldblatt (1973). Plants small to medium, reaching to 55 cm high, usually branched. Corm 1.5-2 cm in diameter; tunics of dark brown fairly coarse fibers often extending upwards in a neck. Prophylls membranous, becoming dry from above. Leaf soli- tary, terete or linear and canaliculate, inserted near the base or the lower part of the stem, and much exceeding the inflorescence. Bract leaves 1-4, 2.5-6 cm long, herbaceous or becoming dry and brown. Spathes herbaceous with dry, light brown margins, attenuate; inner spathe 4-6 cm long, the outer ca. 1 cm shorter. Flowers blue violet with orange yellow nectar guides; outer tepals 2-3 em long, lanceolate, the limb to 1.5 cm long and 1.2 cm wide, spreading to re- flexed at 45°; inner tepals 1.5-2.4 cm long, linear-lanceolate. Filaments ca. 6 mm long, joined in the lower half; anthers ca. 6 mm long. Style branches ca. 1 cm long, the crests to 0.5 cm long. Capsule ovoid, to 1.2 cm long; seeds small, angled. Chromosome number 2n — 12, 24 (South African collections only). 260 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 20 ==] pe | | | | | | | | | @ M. natalensis — I | | | | A M. elliotii | | | \ {hr wu "i | 30', — . "oo 200 200 400 s00 #00 wis I \ | \ — n \ l l | | — I \ |o 200 \ 00 600 900 1000 W _ — — | | | L a Ng | o | 0 , 9 | * | m$ FicunE 4. Distribution of Moraea natalensis and M. elliotii. Flowering time: September to March. Distribution: Eastern South Africa, Swaziland, and Malawi.—Fic. 4. Moraea elliotii is fairly common in the grasslands of South Africa, especially in well-watered highland areas. As defined earlier ( Goldblatt, 1973), it includes a wide range of forms, from the early, spring-blooming eastern Cape plants with a +basal canaliculate leaf, to summer-flowering plants from Natal, Transvaal, and Swaziland, also with a canaliculate leaf, but inserted somewhat above ground level, to terete-leafed, usually late-flowering plants, mainly from the eastern Transvaal. A single collection from Malawi is identical with the latter, and though such a gap in distribution from South Africa to Malawi is unex- pected, the Malawian collection must be assigned to M. elliotii. The species is clearly predominantly South African, and for further details of synonymy, vari- ation and distribution, readers are referred to my earlier treatment of the species ( Goldblatt, 1973). Moraea elliotii is closely allied to species such as M. carsonii from Central Africa and M. polyanthos from the southern Cape, and all three species have a similar flower. It would seem more specialized in its solitary leaf and rela- tively few branches than the multi-leafed and branched M. polyanthos and the 2-leafed M. carsonii. The more reduced M. natalensis, with its short leaf in- serted well above the ground and with a somewhat contracted inflorescence, probably evolved from multi-leafed ancestors via plants like M. elliotii; and M. natalensis is also seen as closely related. MALAWI. CENTRAL REGION: Dedza, summit of Ciwan, Chongoni, Chapman 1176 (SRGH ). 19771 GOLDBLATT—MORAEA 261 | 6. Moraea natalensis Baker, Handb. Irid. 56. 1892. rypes: South Africa, Natal, Sanderson 253 (K, lectotype; S, isolectotype); Sutherland s.n. (K, paratype ).—Fic. 3C. M . erici-rosenii Fries, Wiss. Ergeb. Schwed.-Rhod.-Kongo Exp., 1911-1912, Bot. Untersuch. 1: 234. 1916. type: Zambia, Kalambo, Fries 1345 (UPS, holotype; Z, isotype M. parviflora N.E. Brown, Trans. Roy. Soc. S. Africa 17: 346. 1929. TYPE: South Africa, Transvaal, Tomson's lei Nylstroom, Pole Evans 19668 (K, holotype; PRE, isotype). Plants 15-45 cm high, including the leaf, usually branched. Corm to 1.5 cm in diameter; tunics of dark brown to black fibers. Prophylls membranous with the upper becoming dry from the apex and often lacerated. Leaf inserted well above ground, shortly below the inflorescence, canaliculate to terete, to 20 cm long and shortly exceeding the inflorescence or reduced and about as long as the spathes. Stem with the lowermost internode very long, and the produced leaf inserted in the upper part. Bract leaves if present seldom exceeding 2.5 cm and usually dry and light brown. Spathes herbaceous, becoming dry above, the margins dry and pale brown, the apices attenuate; inner spathe 2-3.5 cm long, the outer ca. 1 cm shorter. Flowers blue mauve with yellow nectar guides; outer tepals 1.4-2 cm long, lanceolate, the limb 0.7-1.4 cm long, up to 1.0 cm wide, re- flexed to 45°; inner tepals to 1.5 cm long, reflexed. Filaments ca. 5 mm long, united in the lower half; anthers 4-5 mm long. Style branches ca. 6 mm long, the crests to 5 mm long. Capsule ovoid-subglobose, 4.5-10 mm long; seeds small, angular. Chromosome number 2n — 12 Flowering time: November to January ( February ). Distribution: Zambia, Malawi, Rhodesia, Mozambique, South Africa; in moist situations, often in vleis and dambos and in seasonal pools.—Fic. 4 The most striking characteristic of M. natalensis is its leaf insertion which is well above ground level at the top of the long lowermost internode. The leaf itself is relatively short, no more than 20 cm long, but more often smaller and sometimes not exceeding the inflorescence spathes. In some forms, notably in the northern part of its range, in Zambia, the stem above the lower internode is much contracted so that the leaf seems to be inserted almost immediately under the inflorescence. In these forms the leaf is often at its shortest, appear- ing almost bractlike and easily confused with the inflorescence spathes. The type of M. erici-rosenii corresponds to this form. Most other collections from Zambia, and Rhodesia, where the species is very common, resemble the typical South African form with a less contracted stem above a longer leaf. The existence of a whole range of forms from the extremely short-leafed, much-contracted type represented by M. erici-rosenii to the relatively long-leafed and extended- stemmed type make it necessary to reduce M. erici-rosenii to synonymy. Moraea natalensis is one of the more widespread species in the genus and occupies a similar ecological niche throughout its range. It occupies rather moist depressions or vleis and dambos in open grassland and in exposed rocky situa- tions which are seasonally moist, and is also found around rock pools. It oc- curs at moderate altitudes of between 1,000 and 2,000 m, though it is also found near the coast in South Africa. It is most closely related to the predominantly 262 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 South African species M. elliotii, which has a single leaf inserted lower on the stem, usually shortly above the ground. MALAWI. CENTRAL REGION: Near Tamanda Mission, Robson 1093 (BM, K, LISC, PRE, SRGH ). 5 Jackson 2299 (K, MAL, PRE, SRGH). Kasungu National Park, Hall-Martin 1364 (SRGH). NORTHERN REGION: S. Vipya, Lwanjati Peak, Pawek 8905 (SRGH, MO). MOZAMBIQUE. MANICA & SOFALA: 12 km from Vila Pery, Torre & Correia 13180 (LISC). ZAMBEZIA: Mujeba, on gea road, Quelimane district, Faulkner K152 (BR, K, PRE, S). RHODESIA. CENTRAL REGION: Marandellas, Dehn 556 (SRGH); Rattray 829 (SRGH ); Day s.n. (SRGH-2129); Collins 10 (K, SRGH). Rusape, Hopkins s.n. (SRGH-6830); Munch 455 (K, SRGH). Salisbury, Eyles 1896 (K, PRE, SAM), 3737 (SAM, SRGH), 6099 (BOL, K, SRGH); Willoughby s.n. (SRGH ). M. siding Selukwe, Taylor s.n. (SRGH-2625). Near Que Que, Bingham 374 (K, SRGH). Gwelo, Loveridge 540 (K, PRE, SRGH). Dar- es Gordon 193017 (K, SRGH). SOUTHERN RE ear Fort Victoria, Plowes 3154 (K, LISC, PRE, SRGH). wESTERN REGION: 5 Brain s.n. (SRGH). Matobo dis- trict, prs 1998 (K, LISC, MO, PRE, SRGH), 2051 (MO, PRE, SRGH). Matopos, Borle 58 (K); Garley 112 (K, SRGH H); Eyles 1144 (ISRGH). Lochview, Cross 335 (SRGH). ZAÏRE. SHABA: Lumbumbashi (Elisabethville), Hock s.n. (BR). Near Mukumbi, Hoff- mann 962 (BR ZAMBIA. CENTRAL REGION: Lusaka, King s.n. (K); Best 4, 10 (K), 82 (SRGH). NORTH- ERN REGION: Mbala (Abercorn) district, Kalambo Falls, Richards Were (K). 25 km W of Kasama, Robinson s.n. (K). SOUTHERN REGION: Machipapa, Mazabuka district, White 6253 (K). Katomo, Patoka Plateau, Sykes 267 (K). Kaloma, Fanshawe 9179 (SRGH); Mitchell 17/29 (SRGH). Between Chomo and Monze, van ee 3084 (BM, K, SRGH). Chomo, Astle 1841 (SRGH); Lawton 1185 (K, SRGH). kaa REGION: Kalenda dambo, Mwinilunga district, ge Redhead 3597 (BR, K, PRE). Zambezi Rapids, Mwinilunga district, Richards 17146 (K, SRGH); Le 178 6224 (K, MO). Kitwe, dambo, Linley 35 (MO, SRGH). Nchanga, Ferrar s.n. ' (SRGH- 480( — 7. Moraea thomsonii Baker, Handb. Irid. 57. 1892; Bot. Mag. tab. 7976. 1904. TYPE: Tanzania, plateau north of Lake Nyassa, Oct. 1880, Thomson s.n. (K, holotype ).—Fic. 5 M. stricta Baker, Vierteljahrsschr. Naturf. Ges. Zürich 49: 178. 1904. type: South Africa, Transvaal, Shilouvane, Junod 563 (Z, holotype; K, LD, isotypes M. tellinii Chiov., Ann. Bot. (Rome) 9: 138. 1911. TYPE: Ethiopi ia, Begemdir & Simen, Debarek, Chiovenda 3007 (F, lectotype); several other oe cited, all from Ethiopia. M. curtisae Foster, Contr. Gray Herb. 127: 46. 1939. type: Kenya, Norok, Noyrosera, 60 km SE of eid Curtis 676 ( GH, holotype). M. trita N. E. Brown, Trans. Roy. Soc. S. Africa 17: 347. 1929. rype: South Africa, Trans- vaal, Lydenburg, bp at b holotype; P, PRE, isotypes). M. parva N. E. Brown, Trans S 8. Africa 17: 347. 1929. TYPE: South Africa, Trans- vaal, Woodbush, Moss 15 5 io pan tes M. mossii N. E. Brown, Trans. Roy. Soc. S. Africa 17: 347. 1929. rype: South Africa, Trans- vaal, B ei ie Moss 15805 (K, 1 PRE, isotype) — Plants small, 12-30 cm high, branched, leafless when in flower. Corm 1-2 cm in diameter; tunics dark brown, of tough reticulate fibers. Prophylls dry and ir- regularly broken. Leaf solitary, quite dry, or lacking at flowering time, occasion- ally a new leaf emergent and then not attached to the flowering stem, slender, terete, long and trailing. Branches short to +sessile, subtended by dry bract leaves. Spathes usually quite dry and papery at flowering, acuminate, becoming lacerated with age; inner spathe 2.5-4 cm long, the outer 5-10 mm shorter. Flower pale blue lilac with yellow orange nectar guides; outer tepals 1.5-2 cm long, lanceolate, the limb to 1 cm, spreading; inner tepals 1.4-1.7 cm long, erect, becoming outspread. Filaments 4-5 mm long, free in the upper third; anthers 1977] GOLDBLATT—MORAEA 963 N WAS ay Ficure 5. Morphology and distribution of Moraea thomsonii (0.5). 4-5 mm long. Style branches 7 mm long, the crest vestigial or 3-5 mm long. Cap- sule ovoid, ca. 1 cm long; seeds small, angular. Chromosome number 2n = 12, 24, 36, 48. Flowering time: (August) September to November (December) south of the equator, December to June (August) north of the equator, in the dry season before the rains begin. 264 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Distribution: Widespread, in dry grassland in the eastern half of Africa; ex- tending from the summer rainfall areas of South Africa through Mozambique, Rhodesia, Malawi, and eastern Zambia to Tanzania, Uganda, Kenya and Ethiopia. Fi. 5 Several species are reduced here to synonymy in M. thomsonii, notably M. tellinii from Ethiopia, M. curtisiae from Kenya, and M. stricta from South Af- rica. These all differ to a small extent from M. thomsonii, but they have its char- acteristic growth habit and peculiar terete leaf, characters which are considered more significant taxonomically than the minor floral differences that are perhaps to be expected in such a widely distributed species. The typical form of M. thomsonii which occurs in Malawi and adjacent Zam- bia and Tanzania has short, almost vestigial style crests, but has relatively broa tepals and style branches. Plants occurring further to the north and also to the south have well-developed style crests. The northern forms, however, have the broad tepals and style branches of the type, while the South African form, earlier treated as M. stricta by me (Goldblatt, 1973), has rather narrow tepals and style branches. Plants occurring in Rhodesia are intermediate between the typical and southern forms and usually have style crests. Moraea thomsonii is one of the more widespread species in the genus, extend- ing almost the length of the eastern half of Africa, from the eastern Cape Province of South Africa to northern Ethiopia. It occurs in open grassland and typically flowers towards the end of the dry season. The leaf and flowering stem are pro- duced at different times, the long, terete leaf emerging after the first rains, and attaining full size during the wet season. The leaf dies back in the dry season, often becoming broken and decayed by the time the stem and, subsequently, the flowers are produced. Owing to the widespread practice in Africa of burning grasslands at the end of the dry season, the leaf of M. thomsonii is often charred or is entirely destroyed, so that flowering specimens frequently lack leaves. Moraea thomsonii is generally easy to recognize in the herbarium because specimens usually lack leaves, or if these are present, it is obvious that the leaf was dead when collected. Difficulty is sometimes experienced in deciding if the leaf was in fact dry when gathered and then the terete nature of the leaf, the short, often sessile branches and dry, almost transparent spathes are sufficient for determination. ErHiOPIA. arusi: Mt. Chillalo, Scott s.n. (K). 1 5 SIMEN: Debarek, Amhara, Chiovenda 3007 (F). ERITREA: Hamasen, Fi iori 895 (F). el pre err Pappi 9 (F). Amba-Dero, Pappi 3541 (F). Belesa, A RA "Terrac no e Pappi 343 (F). At Taclesan, Terraciano & Pappi 415 (F). HaRAR: Mt. Achim, Bally 10054 1 K). SIDAMO: 15 km SE of Meghelli, oe 1827 (F, K). Mogada forest, Mooney 5469 (K). Wadera, emsa 5629 (EAH, F, K). 15 km S of Adola, Bally 3137 (K). Agheremeriam, Gillett 14549 (BR, EAH, F, K). ene Neghelli and Filtu, de Wilde 6675 (WAG). CEN COAST PROVINCE: District around Nyora, Routledge 1908 (K). CENTRAL PROVINCE: Nairobi National Park, Verdcourt 3284 (K). Between Ngong and Kikuyu, van Someren 1437 (EAH, K). Nanyuki, Watt s.n. (K). RIFT VALLEY PROVINCE: Elgeyo, Bat- tiscombe 1184 (EAH, K). Kaptagat, Agnew & Agnew 9031 (MO); pum 2148 (EAH, 4 E). Kapiyet, Dauglish 82 (K). Trans Nzoia, Tweedie 443 (K). Elgon, Lugard 548 (K). Kitale, Thorold 2752 (K). NORTHERN FRONTIER: Near Kisima, iod 8544 (K). NYANZA PROVINCE: Tinderet, Poviano s.n. OUTHERN PROVINCE: Near Kongoni River, Fries & Fries 1538 (K, S. UPS). 60 km SE of Narok. Noyrosera, Curtis 676 (GH). 1977] GOLDBLATT—MORAEA 965 ALAWI. NORTHERN REGION: Nyika oe Park, Pawek 1390 (SRGH). Nyika Pla- teau, Robson 231 (K, LISS, SRGH). CENTRAL PROVINCE: Between Dedza and Ncheu, Jackson 148 (K). Clintembwe, Kota Kota diit Brass 17579 (K, MO). soUTHERN REGION: Mlanje Mt, Burtt Davy 21981 (K); Pawek 3795 (K). Zomba Mt. Salubeni 89 (MAL, SRGH). Mt. Malosa, Whyte s.n. (K). MOZAMBIQUE. MANICA & SOFALA: Bäruè, Serre de Choa, Mendonça 301 (LISC). Be- tween Skeleton Pass and Chimanimani Plateau, Grosvenor 225 (LISC, SRGH). NIASSA: hga near Vila Cabral, Mendonça 776 (LISC). ESIA, EASTERN REGION: Chimanimani Mts., Munch 325, 326 (SRGH). The Cor- ner, Chipset Mts., Wild 3350 (K, SRGH); Sturgeon s.n. (SRGH-30671, K, LISC, PRE); Chase 2973 (BM, MO, SRGH). Melsetter, Plowes 2460 (SRGH), 2800 (LISC, SRGH). Banti North, Umtali, Chase 7828 (SRGH). Inyanga, Leach 8139 (K, SRGH ). Mare River, Inyanga, Wild 3859 (K, MO, SRGH). Slopes of Rukotso, Inyanga, Phipps 734 (SRGH). S of Pungwe View, Methuen 309 (K). SUDAN. EQUATORIA: (reported as Uganda) Imatong Mts., Eggeling 3558 (K TANZANIA. CENTRAL PROVINCE: 1 Mpwapwa district, Gane 133 (EAH, K). EASTERN PROVINCE: Morogoro, Schlieben 1237 , K). NORTHERN PNE Dem Tanner d (K). Liliondo, Masailand, 178 20 (K); Gibbins s.n. (K). slopes of Kiliaman— ro, Greenway 6704 (K, PRE). Hanang Mt., Burtt 4010 (K); LL a 1313 (K ed Mpo- lolo. Moshi district, Haarer 1478 (K). O Oldeani 5 g 2 (K). SOUTHE HIGH- LANDS: Ruhudje-Lupembe, Schlieben 1237 (K, LISC, \ PRE). 185 district, eae 3429 (EAH, K). Mbeya Mt., Geilinger 2797 ( K,Z). WESTERN PROVINCE: Her, Kasulu dis- trict, Eggeling 6195 (BR, EAH, K). S of Sisaga, Kigoma district, Jefford & Newbold 1770 (K). Keto Mt., 15 km from Zambia border, Richards 6186 ( K GANDA. Without precise locality, Bally 10748 (K). NORTHERN PROVINCE: Mt. Debasien, Pegang 2693 ( K = REGION: Lundazi, Nvika Plateau, Pawek 2854 (K, MAL). NORTHERN REGION: Lomi niis Kawimbi, Richards 6116 (K). Subgenus GRANDIFLORA Goldbl. 8. Moraea spathulata (L.f.) Klatt in Th. Durand & Schinz, Consp. Fl. Afr. 5: 152. 1895.—Fic. 7A Iris spathulata B Suppl. Pl. 99. 1781. Type: South Africa, Cape, Langkloof, Wolwekraal, Thunberg s.n. (Herb. Thunberg 1172, UPS, holotype). I. spathacea Thun n Diss. Iride no. 23. 1782. type: as for I. spathulata L.f. s spathacea ( Thunb. ) Ker, Bot. Mag. tab. 1103. 1808, non Thunb., 1787. 1. longi: spatha Klatt, Linnaea 34: 560. 1866. TYPE: South Africa, Cape: Transkei, banks of River (“Tambikuland”), Ecklon & Zeyher Irid. 3 (MO, lectotype). M. spathulata — din at sa Goldbl., Ann. Missouri Bot. Gard. 60: 253. 1973. TYPE: So vaal, near Sabie, Goldblatt 610 (BOL, holotype; MO, isotype ). M. 5 sasi saxosa Goldbl., Ann. Missouri Bot. Gard. 60: 254. 1973. Type: South vies Transvaal, summit of Lans Tom Pass, Goldblatt 612 (BOL, holotype; PRE, — M. Wow der subsp. autumnalis Goldbl., Ann. Missouri Bot. Gard. 254. 1973. TYPE: South Africa, Cape, Transkei , Nyameni Mouth, Port rise ete Strey 8619 (PRE, holotype; NH, isotype ). Plants large, 50-90 cm high, solitary or in small clumps. Corm 1.5-2 cm in diameter; tunics of brown, finely reticulated fibers. Prophylls prominent, brown to pale, firm in texture, brittle, dry, entire or irregularly broken, or frayed at the apex. Leaf solitary, flat or canaliculate, to 1.5 cm wide. Stem simple or occa- sionally bearing one branch. Bract leaves 2-3, often dry and brown, to 15 cm long, rarely overlapping. Spathes herbaceous, or becoming dry and brown from the apex, attenuate; inner spathe 10-14 cm long, outer ca. * the inner. Flower pale yellow; outer tepals 3.5-5.5 cm long, the limb 2-3.5 cm, spreading; inner tepals 3-4 cm long, erect. Filaments 8-12 mm long, free in the upper third; 266 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 10 + | | | | | | r | 20 | | | í+— — | | \ \ \ \ \ | | \ \ \ T E \ E Top ^ O 100200 300 400 500 800 MILES \ —— hq —— o 200 400 $00 $00 1000 KM —— E es ——— | 2 i | L * ) v \ s ome 1-1965 Fıcure 6. Distribution of Moraea spathulata. anthers 8-12 mm long. Ovary 2-3 cm long; style branches 1.2-1.8 cm long, the crests to 1 cm long. Capsule 3.5-5.5 cm long; seeds large, flattened. Chromosome number 2n = 12 (South African plants only ). Flowering time: (November) December to March (early April). Distribution: Open grassland in the Inyanga and Chimanimani highlands of Rhodesia and Mozambique, also in South Africa, Lesotho, and Swaziland.—Fic. 6 The tall, summer-flowering Moraea frequently collected in the Chimanimani and Inyanga highlands of Rhodesia and Mozambique is clearly closely allied to the M. spathulata complex of South Africa. The plants from the Chimanimanis in particular are virtually identical to the Transvaal and Swaziland forms which I previously referred to M. spathulata subsp. saxosa Goldbl. (Goldblatt, 1973). Since 1973 when I recognized M. spathulata as comprising four subspecies, I have had the opportunity to observe living plants in several places and to see more herbarium material. As a result, it has become clear that my attempt to sub- divide M. spathulata was not satisfactory, and the proposed subspecies did not accurately reflect the variation found in the species. The subspecies autumnalis from the Transkei is merely a very early blooming coastal form and must be in- cluded in the typical form. Moraea spathulata in the southern part of its range in the Knysna district of the Cape Province blooms from July to September but not unusually in June or even May, thus the March to May blooming subsp. autumnalis is not particularly unusual. The remaining two subspecies, subsp. transvaalensis and subsp. saxosa, both from the Transvaal and Swaziland, dif- fer from one another mainly in that the lower-altitude subsp. transvaalensis 1977] GOLDBLATT—MORAEA 967 — Ficung 7. Moraea species. -A. M. spathulata—B. M. inyangani—C. M. schimperi (0.5). 268 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 forms clumps, while the higher, altitude subsp. saxosa appears to be predomi- nantly solitary in habit and has a slightly longer ovary and capsule. However, when the entire pattern of variation in M. spathulata is considered, both the tendency to solitary habit, and the longer ovary of subsp. saxosa and the features subsp. transvaalensis now seem altogether too insignificant to make any taxonomic recognition worthwhile. It seems far more useful to recognize M. spathulata as a single, complex, and variable entity. North of the Limpopo River M. spathulata thus extends as far as the Inyanga region of Rhodesia-Mozambique. The largest forms occur in Inyanga, some plants even bearing branches while rather dwarfed forms occur in the Chima- nimanis. Differences between the two appear significant until the few collections from the intervening highland areas in Mozambique are considered, and these prove intermediate in all characteristics. MOZAMBIQUE. MANICA & sOFALA: Tsetserra, Exell, Mendonga & Wild 332 (LISC); Torre & pasha 15687 (LISC). Gorongosa Mts., summit, Tinley 2436 (SRGH). Serra Mes- so ar orre & Correia n (LISC). Between Mussapa River and frontier = Tendera, e d» eh 13159 (LISC). Bárué, Serra de Choa, Torre & Correia 15485 RU. EASTERN REGION: Melsetter, Hanmer s.n. (SRGH-18354); a 1421 (BM, K, SRGH). Melsetter district, d farm, Crook 318 (SRGH). Glendingwe Estate, Plowes 3477 (SRGH). Pork Pie Hill near Melsetter, Bamps, Symoens & van den Berghen 776 (BR, SRGH). Inyanga, Chase 4351 (BM, K). Inyangani Mt., Chase 8124 (BM, K, LISC, PRE, SRGH); Plowes 2429 (SRGH 1 Whellan & Davies 984 (K, SRGH); Fries, Nrolindh Ç Weimarck 4967 (BM, BR, PRE). Mt. Mov opus Stapleford, 1 8 5711 (K, MO, PRE, RGH). Bant North, Umtali district, Wild 4522 (K, LISC, MO, PRE, SRGH). Himalayas- Engwa, Wild 4456 (K, LISC, SRGH). a Mts. near Umtali, Obermeyer 2113 (P 9. Moraea schimperi (Hochst.) Pic.-Serm, Webbia 7: 349. 1950.—F'c. 7C. Hymenostigma schimperi Hochst., Flora 27: 24. 1844. Type: Ethiopia, Begemdir & Simen :chedcap," Schimper 1173 (B, holotype; BM, F, K, M, MO, P, S, isotypes). H. tridentatum Hochst., Flora 27: 25. 1844. type: Ethiopia, Begemdir & Simen, Barnam, achit, Schimper 1296 (K, lectotype ). Vieusseuxia schimperi ( Hochst.) A. Rich., Tent. Fl. Abyss. 2: 305. 1850 (Hist. Nat. Bot. V. . (Hochst. ) - Rich., Tent. Fl. Abyss. 2: 305. 1850. Iris diversifolia Steud., m.s. ( cf. A. Rich., Tent. Fl. Abyss. 2: 305. ) ido diversifolium 'Steud. ex Klatt, Linnaea 34: 572. 1866, nom. erus superfl. TYPE: as for menostigma schimperi ipe d e ex ven Baker, J. hs Soc., Bot. 16: 130. 1877. M. welwitschii Baker, Trans. Linn. Soc. London, Bot., Ser. 2, 1: 5 1878. rype: Angola, Huh. Lopollo River, 5 1548 (BM hulntepe: K, , P, isotypes). M. zambeziaca Baker, . Afr. 7: 340. 1898. TYPE: 29 Mangaja hills, Meller s.n. (K, holotype ) M. hockii De Wi Spec. Nov. Regni Veg. 11: 540. 1913. Type: Zaire, Shaba, between Buggege d Tokoni, Hock s.n. (BR, holotype). Plants 2040 cm high, unbranched. Corm 1.5-2 cm in diameter; tunics brown, fibrous. Prophylls 3-5, large and prominent, entire, firm, usually dark brown. Leaf solitary, basal, canaliculate to flattened, 9.6-15 mm wide, emerging as flowering begins, but eventually much exceeding the inflorescence. Stem usually short at flowering time but elongating in fruit. Bract leaf usually solitary, 10-15 cm long, becoming dry and brown. Spathes usually dry and brown, (6-)7-10 (-12) cm long; outer spathe about %4 the length of the inner. Flower blue purple with yellow white nectar guides; outer tepals lanceolate, 4—6.5 cm long, the limb 1977] GOLDBLATT—MORAEA 269 equal to or slightly exceeding the claw; inner tepals erect, 3.5-4.5 cm long. Fila- ments 9-15 mm long, united in the lower half; anthers 8-12 mm long. Ovary 1.5-2 cm long; style branches 1.5-2 cm long, the crests 1-2 cm long. Capsule 2.5-3.5 cm long; seeds flattened and +discoid. Chromosome number 2n = 12. Flowering time: From the end of the dry season into the early wet season, August to November (December) south of the equator; November to May, oc- casionally until July in West Africa and Ethiopia. Distribution: Widespread from Rhodesia north and west through Central and East Africa, to Angola, Zambia, Malawi, Mozambique, Tanzania, Zaire, Burundi, Sudan, Ethiopia, Cameroons and Nigeria; often in marshy or wet situ- ations, occasionally in grassland.—Fic. 8 Moraea schimperi has been given a surprising number of names which seems remarkable in the light of its singular morphology and habit. This is probably explained more by its great geographic range than by any morphological dif- ferences, and in fact most of its synonyms have been applied to plants from different areas of Africa. Moraea schimperi extends from the Rhodesian-Mozam- bique highlands in the south, west to Angola and north, through Zaire and Burundi to southern Sudan and Ethiopia. It also occurs in the higher areas of Nigeria and Cameroons, with a large break in the range over the Congo Basin. Significantly it is also absent from Uganda and Kenya, where the highlands may ve of too recent origin for it to have become established Moraea schimperi usually occurs near a permanent water source, either un- derground or at the surface, in a vlei or dambo, or along a stream, but is also found in well-drained grassland in areas of high rainfall. It flowers at the end of the dry season and continues in bloom in the early wet season. Thus in the southern tropics it blooms from September through October and into November and early December. On the equator and to the north, plants most often begin flowering in December to February, and continue into March and May. The morphology of M. schimperi is most distinctive. The flowering stem appears be- fore or as the new season's leaf emerges, and early in the season the bright blue flowers are borne well above the leaf apex. The prophylls, bracts, and spathes are large, and their usually rich brown color is alone sufficiently characteristic to permit determination. As the season progresses, stem and leaf continue to grow, and by the time the seeds are ripe the stem may be a meter high and the leaf much longer. There is only a small degree of variation from this pattern, notably in plant size, with forms from Nigeria-Cameroons and also some in Ethiopia being more slender than the comparatively robust plants in the southern part of its range. hough clearly related to the M. spathulata complex, as can be seen from the similarity of the bracts and prophylls, M. schimperi is unusual in this al- liance in having blue purple flowers. The treatment of M. schimperi here differs somewhat from that of Geerinck (1970) in that M. arnoldiana is not con- sidered a synonym. The latter is a late-blooming species from Zaire and is as- signed to synonymy under M. macrantha in this treatment. ANGOLA. BIE: Chitembo, Barbosa & Moreno 12245 (LISC). cuANzA SUL: Murta d» Silva 857 (COI, LISC). nuAMnBo: Rio Cuanza, Mucosa, Monteiro d Murta 1884 (COI, 970 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 — — | ETIT * 0 yoo 200 300 400 500 800 MILES — — | — — (0/90 C« ASo|ào 0 200 400 $00 800 1000 KM = oes 1— ss — — | — \ m \ \ | — oj — FicunE 8. Distribution of Moraea schimperi. LISC, SRGH). Huambo (Nova Lisboa), da Silva 3188 (LISC, PRE). HuiLA: Missae Cato- lica, Huila, Santos 62 (LISC). Lopollo River, Welwitsch 1548 (BM, be K, LISU, P Near Vila Arture de Paiva, Leach d» Cannell 13861 (K, LISC). Lubango (Sa de Baneira), Rio Caia, Henriques & Brites 1137 (K, LISC, LISU, PRE). Humpata, Mendes 1816 (LISC). Cheila Mts., Gosseiler 13316, 13317 (LISC). MALANJE: Gossweiler 922 RUNDI. Bururi, Becquet 162 (BR, EAH, K, WAG); Reekmans 1041 (BR, EAH). Near ei Lewi ille 6131 (BR, MO). ooN. WEST: Bauer 22 (K). Bamenda, 4 km NE of Bambili, Bauer 154 (K). E i pone nda and Santa, Keay s.n. ( FHI-28348, K, P). East: Nganha Mts., E. Ngoaun- 1977] GOLDBLATT—MORAEA 9 271 déré, Jacques-Felix 8632 (P); nd Wilde 4521 (WAG). E of „ Raynal S Raynal 12236 (P); de Wilde 4312 (WAG). Dschang, Mt. Bamboutos, Villiers 672 (P); Jacques- Felix 2707 (P); Saxer 88 (K, m Z). Dschang, Meurillon 765 (B B Djuttitsa, Meuril- lon 179 (BR), 745 (P), 1212 (BR, K, P). Mt. Santa, Jacques-Felix 2810 (P). Gotel Mts., NNE Banyo, Letouzey $592 (P). EruiopiA, ARUSI: Between Shashamane and Dodollo, de Wilde 6823 (WAG). Near Asela, de Wilde 6595 (MO, WAG). Near Kofole, Mooney 7106 (F, K, S). Between Bekoji and Adaba, Gilbert 1150A (K). BEGEMDIR & 1 a Michubbi, Pichi-Sermolli 2674 (F, K). "Enschedcap," d 1173 (B, BM, F, K, MO, P, S). Sering, Schimper 1536, 1537 (P). Hanan: Gar Mullata Mts., Gillett 5332 mI F, K, P, S); Burger 1706 (K), 2933 (F, K); de Wilde 6335 (WAG). sHoa: Negri 413 (F). Between Addis Ababa and the Blue Nile, 1 957 (F). Mt. Zuguola, Buscalioni 2158 (F). sipaMo: S of Agere Selam, de Wilde 6 i721 (WAG). SE of Agere Selam, de Wilde 10298 (MO, PRE, WAG). WOoLEGA: Bari, 1 79 Sats N of Beica, Mooney 7 7729 (EAH, K). MALAWI. NORTHERN REGION: Chelinda Bridge, Nyika, Pawek 1407 (SRGH). Nyika Plateau, Cottrell 3, "65 (SRGH); Robson 326 BR, K, LISC, SRGH). Mzuzu, Mary- mount, Pawek 5833 (MO, SRGH). Vipya Mts., Chapman 1702 (K, LISC, SRGH); cay (SRGH). CENTRAL PROVINCE: Dedza, Brass 17632 (K, MO, SRGH); Kulumule 2 (K, RGH). Chongoni Forest Reserve, Dedza district, Salubeni 25 (MAL). SOUTHERN PROVINCE: wa nÀ Plateau, Mt. Mulanje, Newman & Whitmore 653 (BR, K, SRGH, WAG). Mt. Mulanje, Forbes 67 (EAH); imum, 6306 (EAH, PRE). 1 Plateau, Mulanje Mts., Pawek 3793 (MAL). Limbe, off Cholo road, Moriarty 399 (MAL). MOZAMBIQUE. NIASSA: Amayaniba near Mandimba, M 666 (LISC). TETE: Between Furancungo a A onia, Torre 3362 (LISC). zAMBEziA: Namuli hills, Last s.n. (K). Near Gurué, Torre d» COE 15923 (LISC). Serra m Gurué, Mendonça 2229 (LISC). NIGERIA. NORTHERN REGION: Vom, Jos Plateau, Dent Young 245 (K); McClintock 199 (K). Mambila Plateau, Tuley 1929 (K); Latilo & cas = SHE 34380, BR, K, S, WAG); Wit, Gbile & Daramola 2076 (FHI); Hepper 1774 R, k, P); Gbile & Daramola s.n. (FHI- 62880, Ye: G). Between Nguroje and Maisamari, c 2640 (FHI). Vogel Peak, Hepper 1504 ( K, P). RHODESIA. EASTERN REGION: Melsetter, Markhams Kloof, Crook M151 85 LISC, MO, PRE, SRCH). Inyanga downs, Drewe 18 (SRGH). CENTRAL REGION: Salisbury, Eyles 1791 (K, PRE, SAM, SRGH). 5 Dam, Wild 3871 (K, SRGH ). Manda Rattray 562, 309 (BM, SRCH); Corby 196 ). DA? ORIA : d 19 55 Andrews 1895 (K); Johnston 1492 (K); MacDonald SO (BM). Kippi, "Chipp $9 (K). TANZANIA. SOUTHERN HIGHLANDS: Poroto Mts., Geilinger 2611 (K, Z); McGregor 3 (K EAH). Between Chunya and Mbeya, Burtt 6225 ( BM, BR, EAH, F, K). Chunya cscam- ment, Richards 25795 (K). Njombe district, Richards 6585 (K); Gillett 17781 (EAH, K). Itaka, Greenway 3651 (EAH, K, PRE). Mufindi, Paget-Wilkes wie (EAH, MO). 2E Plateau, Richards 18460 ( K, SRGH ); Davies s.n. ( K). SOUTHERN PROVINCE: 8 E W of Songea, Milne-Redhead & Taylor 8377 (K). NORTHERN PROVINCE: Between Dongobesh and Mkulu, Haarer 11613 (K). WESTERN PROVINCE: Sumbawanga, Van Rensburg s H). ZAÏRE. SHABA (Katanga): Gare de Biano, Schmitz 4893 (BR. VAG). Kipopo, Schmitz 5495 (BR). Parc Nationale de l'Upemba, Robyns 3940 (BR). 0 Hen S of Jadot- = — ~ ville, — 4055 (BR). ug Kendo, de Witte 537 (BR, WAG, AN NORTHERN REG Mbala (Abercorn) district, Lumi River I Richards 5844 ( BR, K, SRGH). Bu rl Mbala district, Richards 13138 (K, SRGH). Mansa a meo. Fries 608 (UPS). ke Shiwa Ngandu, Greenway & Praga 5709 ne H). CENT REG p Pw nje, Fanshawe 6727 (SRGH). E. ASTERN REGION: Nyika Platea 3 3012 (K E, SRGH). WESTERN REGION: of Kakema River, n district, Milne- ee 5 (PRE). 10. Moraea muddii N. E. Brown, Trans. Roy. Soc. S. Africa 27: 346. 1929. TYPE: South Africa, Transvaal, Mac Mac Creek, Sabie district, Mudd s.n. (K, holotype). Plants 15-40(-70) cm high, solitary. Corm ca. 1.5 cm in diameter; tunics fine, usually pale. Prophylls, short, dry, pale brown, usually irregularly broken. Leaf linear, canaliculate ca. 5 mm wide, exceeding the inflorescence. Stem un- 979 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 branched. Bract leaves 2-4, herbaceous, 8-12 cm long. Spathes herbaceous; inner spathe 7-12 cm long, the outer ca. % the inner. Flowers yellow; outer tepals lanceolate, 3.5-5 cm long, the limb 2-3 cm long; inner tepals to 3.5 cm long, erect. Filaments to 1 cm long, free in the upper third; anthers 8-11 mm long. Ovary ca. 1.5 cm long; style branches to 1.3 cm long, the crests to 1 cm long. Capsule ca. 2 cm long; seeds flattened and disc shaped. Chromosome number not known. Flowering time: Late August to November. Distribution: Eastern mountains of southern Africa, from the eastern part of the Cape Province to the Chimanimani Mountains of Rhodesia and Mozam- bique; usually in wet situations in grassland.—FIc. The few collections of a rather short, large yellow-flowered Moraea from the Chimanimani Mountains of Rhodesia and Mozambique match very closely M. muddii, a species previously recorded only from South Africa. The Chimani- mani plants resemble M. muddi not only in form, but like it, are early bloom- ing, and generally occur in moist habitats. The range of M. muddii extends from the Chimanimanis south, along the eastern escarpment of South Africa to the Hogsback in the eastern Cape. The Rhodesia-Mozambique plants differ mainly in their rather early flowering, from August to September, in contrast to October to November in South Africa, but otherwise correspond very closely. Moraea muddii is related to the M. spathulata complex, particularly to soli- tary-growing species such as M. moggii, M. hiemalis, and M. carnea, amongst others, and is distinguished by its shorter size, often smaller flower, and early blooming habit. MOZAMBIQUE. MANICA & SOFALA: Near summit of Chimanimani Mts., Grosvenor 195 (LISC, SRGH, UPS, WAG). RHODESIA. EASTERN REGION: Chimanimani Mts., Garley 177 (SRGH); Loveridge A42 (SRGH). Bank of Bundi River, Melsetter district, Whellan 2153 (SRGH). 11. Moraea inyangani Goldbl., sp. nov. Type: Rhodesia, vlei on Mt. Inyangani, 8,000 ft, Wild 5519 (SRGH, holotype; K, LISC, M, PRE, isotypes ).—Fic. 7B. Planta parva, 15-30 cm alta. Tunicae cormi pallidae, tenuissimae. Folium solitarium, NE eue inflorescentium excedens. Caulis simplex ferens bracteis. Spathae herbaceae, interiora 5-8 cm longa, exterior paulo Ba Flores luteis; tepala exteriora ca. 2.5 cm longa, limbo ca. 1.5 cm longo, effuso; interiora -2 cm, erecta. Filamenta 4 mm longa; antherae ca. 6 mm longae. Germen ca. 1.4 cm ees rami styli 8-9 mm longi; cristae 4-8 mm longae Plants small 15-30 cm high. Corm to 1 cm in diameter; tunics of fine, pale fibers. Prophylls brown, papery, irregularly broken. Leaf solitary, canaliculate, appearing terete with the margins tightly inrolled, to 3 mm wide and exceed- ing the inflorescence. Stem unbranched. Bract leaves 3(-4), herbaceous, 5-8 cm long. Spathes herbaceous with dry, brown acute apices; inner spathe 5-8 cm long, the outer only slightly shorter. Flowers pale yellow; outer tepals ca. 2.5 cm long, lanceolate, the limb ca. 1.5 cm long, spreading; inner tepals erect, 1.5-2 cm long. Filaments 4 mm long, free in the upper half; anthers ca. 6 mm 1977] GOLDBLATT—MORAEA 273 20 — A M. muddii W M. inyangani — Ro e M. angolensis | | | 4 l T 30. — —— pu L px 0 100200 300 400 300 600 MILES ———— O 100 400 800 $00 1000 KM 20 ho 0 \ 10 20 30 40 30 Em FicurE 9. Distribution of Moraea muddii, M. inyangani, and M. angolensis. long. Ovary ca. 1.4 cm long; style branches 8-9 mm long, the crests 4-8 mm long. Capsule and seeds unknown. Chromosome number not known. Flowering time: September to October (also in April). Distribution: Rhodesia, endemic on Mount Inyangani, Inyanga district, in moist situations at high altitudes.— FIG. 9 The distive, small-flowered Moraea inyangani appears to be a very local spe- cies, having been recorded only from the higher altitudes of Mount Inyangani, in the Inyanga Highlands of Rhodesia. It is most closely related to M. muddii, which occurs to the south on the Chimanimani mountains of Rhodesia and Mozambique and in South Africa. Moraea muddii has a larger flower and a canaliculate leaf in contrast to the subterete leaf of M. inyangani. Both species appear to occupy the same habitat, high altitude grassland in moist situations, and both flower early in spring. RHODESIA. EASTERN REGION: Inyanga district, Mt. Inyangani, Leech 8140 (SRGH); Corby 808 M SRGH). Summit ridge of Inyangani, damp flush, Drummond d» Robson 5830 (K, PRE, SRGH). Inyangani, wet flush below summit, nus res Ace. Vlei on Mt. Inyangani, Wild 5519 (K, LISC, M, PRE, SRGH ); Burrows 570 (SR 12. Moraea angolensis Goldbl., sp. nov. tyre: Angola, Moxico, Rio Culiti, Capello & Ivens 18 (LISU, holotype). Planta graciles. Folium SS Caulis 35-40 cm alta, ug im tria, illa 4-4.5 cm longa. Spatha 4-5 cm longa. Flores parvi; tepala exteriora ca. 3.5 cm longa, ungues ca. 2 cm; tepala interiora ca. 3 cm lon 17 Filamenta pe 11 mm longa, libera a apicem r mm; . 6 mm longae. Germen 1-1.2 cm longum; rami styli ca. 8 mm longi; cristae ca. 8 mm longae Plants slender, 35-40 cm tall. Corm unknown. Leaf solitary, basal, ca. 2 mm wide and much exceeding the inflorescence, channeled. Stem erect. Bract leaves 974 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 3, 44.5 cm long with dry brown apices. Spathes 4-5 cm long, herbaceous, with the upper 5 mm dry and brown; outer spathe about % as long as the inner. Flow- ers P yellow; outer tepals ca. 3.5 cm long, the claw 2 cm long; inner tepals ca. 3 cm long. Filaments to 11 mm long, free near the apex for 2 mm; anthers 6 mm long. Ovary 1-1.2 cm long; style branches ca. 8 mm long, the crests to 8 mm long. Capsule and seeds unknown. Chromosome number unknown. Flowering time: August. Distribution: Plains of southeastern Angola.—F'c. 9. It is with some hesitation that I describe this species as it is based on two very poorly preserved specimens. The holotype and only gathering was col- lected by Capello and Ivens in August 1885 on their expedition across Africa from Angola to Mozambique. Dr. I. Melo of the University of Lisbon and Dr. E. Mendes, Director of the Junta de Investigacoes do Ultramar, Lisbon, Portu- gal have kindly aided me in localizing the place of collection, Rio Culiti, as south- eastern Angola between latitude 14°50’S and 15°50’S and longitude 19°20’E and 21°50’ E. No other species of Moraea are known from anywhere near this area. The plants are quite different from any other tropical African species, per- haps most resembling M. muddii and M. inyangani both from the highlands of southeast Africa. The slender form, very short bracts and spathes, combined with a small flower make it easily distinguishable from these. ANGOLA: Moxico, Rio Culiti, Capello & Ivens 18 ( LISU). 13. Moraea bella Harms, Bot. Jahrb. Syst. 28: 364. 1901. Type: Tanzania, Southern Highlands, Uhehe, Goetze 698 (B, holotype).—Fic. 10. Plants solitary, slender, (30-)40-60(—70) cm high, very rarely branched. Corm ca. 1.5 cm in diameter; tunics of pale medium to coarse reticulate fibers. Prophylls small, light brown, dry and irregularly broken. Leaf solitary, canalic- ulate, 2.5-6 mm wide, inserted at ground level, and exceeding the inflorescence. Stem slender. Bract leaves 2-4, widely spaced, 6-7 cm long. Spathes dry and brown in the upper part or entirely so; inner spathe 6-10 cm long, the outer % to % the length of the inner. Flowers pale yellow, with a darker nectar guide and often conspicuously veined and spotted; outer tepals 4.5-5.5(-6.5) cm long, the limb 2.5-4 cm, spreading; inner tepals 3.5-5(-6) cm long, erect. Filaments 9-14 mm long, free in the upper third; anthers 8-10 mm long. Ovary 1.5-2 cm long; style branches 1.3-1.6 cm, the crests 0.8-1.5 cm long. Capsule ovate-oblong, 2-3 cm long; seeds flattened, + triangular to discoid. Chromo- some number not known. Flowering time: (Late February) March to June (July). Distribution: Northern Mozambique, Tanzania, Malawi, Zambia, Zaire; only in seasonally or permanently waterlogged habitats such as marshes, vleis, dambos; flowering towards the end of the wet season and in the dry season.— Fic. 10 Moraea bella can readily be recognized by the combination of several morpho- logical features and its moist habitat and late flowering. It is a slender plant 1977] GOLDBLATT—MORAEA 975 FicurE 10. Morphology and distribution of Moraea bella (0.5). with a long, narrow leaf, and 2-4 comparatively short, widely spaced bract leaves. The yellow flowers, which are sometimes characteristically speckled, appear from late February through the end of the wet season and well into the dry season in July. The peak flowering period for the species is in April. Moraea bella is almost invariably found in very wet situations, usually in dambos but 976 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 20 ö | — f | O M. macrantha I l | | e @ M. verdickii | | | f f 1 L l \ \ " —_+— —ů— 30, pa mm 1 | | 0 100 200 300 400 $00 600 MILES | \ — FT n \ — | | | \ © 200 \400 #00 - 06 | — — ae — | 20 to | 0 | 10 FicunE ll. Distribution of Moraea verdickii and M. macrantha. on occasion simply in damp, poorly drained grassland which retains moisture well after the rains are over. Though described early this century, the name M. bella has been until now overlooked. Plants belonging to this species have either been assigned to M. angusta, a common and quite unrelated Cape species, or were included in other species, as for example by Geerinck (1970) who included specimens of M. bella 1977] GOLDBLATT—MORAEA 977 in his very broad concept of M. textilis. Moraea bella is closely related to M. verdickii, a larger-flowered species with the same general range and to the M. macrantha-ventricosa-textilis complex of south central Africa. MALAWI, NORTHERN REGION: Mrzimba, W 1195 (K), 1339 (BM). 8 bo, Mzimiba district, Pawek 8460 (K, MO, SRGH). Fort Hill, Whyte s.n. (K). ear Fort Hill, Say Evans & Erens 705 (BR, K, an SRGH ). Chelinda bridge, Rumphi io Pawek 219 ). CENTRAL REGION: Mchinji district, Brummitt 10206 (K). Kasungu Game eserve, omi 11603 (K); Hill-Martin 1776 (PRE, SRGH). 22 km S of Kasungu, Moriarty 355 (M d NIASSA: Vila Cabral, Pedro & Pedrogao 3629 (EAH). ANZANIA. SOUTHERN HIGHLANDS: Iringa, Lynes 269 (K); Pedersen 990 (DSM); van Rensburg 677 (K). Between Iringa and Dabava, Eggeling 6122 (BR, EAH, K). Lunzua Ag- PER Station, marsh, Richards 5173 (BR). WESTERN PROVINCE: 65 km $ of Sumbawanga, a i3 Libo 10072 (K, LISC, SRGH). Ufipa district, Isopa, Robertson 641 (K, WAG). SHABA: Near Kapona, Plateau des Muhila, Schmitz 1652 (BR). Marungu- Mcr van den Bromde 1 (BR). Jadotville, vallée de la Mulende, Dubois 1274 (K), 1074 A Kankela Valley, Manika Plateau, Malaisse 8564 ). AMBIA. EASTERN REGION: Chipata ( Fort Jameson) Fanshawe 4501 (EAH, K, SRGH); van Rensburg 2111 (K, SRGH); Munch 459 (K, LISC, SRGH). Lundazi, Fanshawe 9291 (K, SRGH). Lukusuzi Game Reserve, Sayer 205 (SRGH). CENTRAL REGION; Chakwenga headwaters, E of Lusaka, Robinson 6473 (K, SRGH). Kawambura, Fanshawe 3674 (K). tween Undaunda and Rufunsa, Kornas 1519 (K). NORTHERN REGION: N of Mbala ( Aber- corn), Greenway 6219 (K, EAH, PRE). “Fwambo,” Nutt s.n. (K); Carson s.n. (K). Mbala district, Kawimbe road, Richards 21407 (K, MO); Sanane 1225 (K, PRE); Robertson 618 (EAH, K); Richards 897 (K 0, 1367 (K). Kambole escarpment, Richards 9989 (K). Chisinga Ranch near Luwingu, Astle 524 ( K, SRGH ), 3017 (SRGH). — 14. Moraea verdickii De Wild., Ann. Mus. Congo, Sér. 4, Bot. 1: 17. 1902. TYPE: Zaire, Shaba, Lukafu, Verdick 281 (BR, holotype).—Fic. 12A Plants large, 45-75 cm high, unbranched. Corm ca. 2 cm in diameter; tunics coarse, of pale or occasionally dark fibers. Prophylls pale to dark brown, dry, broken and fibrous apically. Leaf linear, shortly exceeding the inflorescence, canaliculate or + flat, 4-12 mm wide, inserted near the base but often up to 10 cm above ground. Stem bearing 2-3(-4) herbaceous bract leaves, usually widely spaced but occasionally overlapping. Spathes herbaceous, with a brown, acute apex; inner spathe 9-15 cm long, the outer ca. % the inner. Flowers yellow; outer tepals (5-)6-7.5(-10) cm long, the limb 3-5 cm long, spreading; inner tepals erect, 4.5-7 cm long. Filaments 11-16 mm long, free in the upper third; anthers (10-)11-14 mm long. Ovary usually exserted, 1.5-2(-2.5) cm long; style branches 1.7-2.5 cm long, the crests 1-1.7 cm long. Capsule and seeds not nown. Chromosome number not known. Flowering time: Late November to February (March). Distribution: Grasslands and open bush, occasionally in damper situations, in eastern Angola, northern and western Zambia, Zaire, Tanzania, Malawi, and central Mozambique.—F'c. 11 Moraea verdickii is one of the taller species of Moraea and has particularly large flowers. It occurs in a belt across south tropical Africa from eastern Angola through southern Zaire and Zambia to Malawi, southwestern Tanzania and Mozambique. Though recognized as early as 1902 by De Wildeman, it has in the past been included in M. ventricosa in herbaria, and most recently was in- 978 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficure 12. Moraea species—A. M. verdickii.—B. M. ventricosa.—C. M. textilis (x0.5) 1977] GOLDBLATT—MORAEA 279 cluded with M. ventricosa in M. textilis by Geerinck (1970) who held a very broad concept of this species. A detailed and critical comparison of a large number of specimens of M. ver- dickii and its close allies, M. ventricosa, M. textilis, and also including the related M. macrantha was made in the course of this study. In my estimation the best solution to the systematics of this group is to recognize all four species. Moraea verdickii can be distinguished from its close allies by its large, consistently yel- low flower, with the outer tepals ranging from (5-)6-10 cm long and the anthers (10-)11-14 mm long; few bract leaves, generally 2 or 3; its habitat, grassland and open bush; and its early flowering, from late November to February, oc- casionally until March. Moraea macrantha has similar large flowers which are always dark blue, has 4-5 bract leaves, and flowers later, from mid February to July. Moraea ventricosa has smaller, blue or white flowers, 3-6 bract leaves, often grows in wet situations, and flowers from February to May; while M. tex- tilis, with either yellow or blue flowers, generally has 5-7 bract leaves, and reaches peak flowering in May. ANGOLA. MOXICO: Lusavo falls, Milne-Redhead 4088 (BR, K). MALAWI. CENTRAL PROVINCE: Dedza, Jeke 79 (K, SRGH). Chongoni forest, Dedza, Banda 526 (SRGH); Salubeni 1253 (K, MAL, SRGH). Dedza mountain area, Adlard &. ed 561 (K, LISC, SRGH). Kirk Range, Ncheu-Neno road, Robson 1404A (K, LISC uc PPW NIASSA: 80 km from Mines de Massangulo to Vila Cabral, Correia 154 Near Mtengula, Seddon 7 (K). rere: 8 km from Mlangeni, on Ncheu-Dedza road, Brummitt 8609 (K, SRGH). Kirk Range, Ncheu- 1 road, Robson 1404 (K). Massangulo, Gor & Sousa pes (K). ZAMBEZIA: Grirué near Vila wu ' Torre 5078 (LIRC). TANZANIA. SOUTHERN HIGHLANDS: Iringa district, 25 km S of Dabaga, Polhill & Paulo 3 (BR, EAH, K, mpl PRE). 50 km N of M ire a, Bally & Carter 16475 (EAH, SRGH ). Ca. 120 km N of Mbeya on Itigi road, Boaler 843 3 SOUTHERN PROVINCE: Matengo hills, Songea, Milne-Redhead & paulo, 9024 ( LISC, PRE, SRGH). ZAIRE. SHABA: SE Lubumbashi ( Elisabethville ), a 3721 (BR). Near Lubumbashi, Hirschberg 5 58 (PRE). Keyberg, Schmitz 1230 (BR). Derubo, Quarré 1008 (BR). Marie José, Quarré 1524 (BR). Lupaku River, Kassner 2466 (BR, K, Z). Ho n Kisinde, Dubois 1074 de WAG). Mitwaba terr. River Kenia, de Witte 2470 (K, SRG MBIA. NORTHERN PROVINCE; Kambole escarpment, Richards ru (K, UPS). Item- bwe Corge, Mbala (Abercorn ) district, Richards 12055 (EAH, K, MO, SRGH). Itembwe , SRGH). Mhala Kalambo falls road, Richards 17080 (K, SRGH); Bullock s.n. (BR, K). Mpika, Fanshawe 1890 (BR, K, SRGH). 40 km N of Mpika, Wil- liamson 1470 (K, SRGH). S of Nchelenga, Kawambwa district, W 7 ond 1233 (SRGH ). Chishinga ranch, Astle 1395 (SRGH). WESTERN PROVINCE: Matonchi farm, Mwinilunga district, Milne-Redhead 3930 (BR, k, LISC, PRE). O e T = S = = a m^ — a — > oo ° eo ~ 15. Moraea macrantha Baker, Fl. Trop. Africa 7: 340. 1898. rype: Malawi, Northern Province, without precise locality, Whyte s.n. (K, holotype). M. arnoldiana De Wild., Ann. Mus. Congo, Sér. 4, Bot. 1: 16. 1902. Type: Zaïre, Shaba, Kasenga, Lukafu, Verdick 606 ( BR, holotype Plants solitary, unbranched, 50-70 cm "€ Corm ca. 1.5 cm in diameter; tunics of pale fine to medium fibers. Prophylls brown, soft textured and ir- regularly broken to fibrous. Leaf solitary, linear, basal, canaliculate, to 7 mm wide, much exceeding the inflorescence. Stem unbranched. Bract leaves 4-5, herbaceous, usually overlapping, attenuate, 6-10 cm long. Spathes herbaceous, with a dry acute apex; inner spathe 9-13 cm long, the outer ca. % the inner. 280 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Flowers blue violet with pale nectar guides; outer tepals (5.7-)6.5-8 cm long, the limb + equal or exceeding the claw, 3.0-4.5 cm long; inner tepals 5.5-7.5 cm long, lanceolate. Filaments 1.3-1.7 cm long, free in the upper half to third; anthers 1.2-1.5 cm long. Ovary ca. 2 cm long, often enclosed in the spathes; style branches ca. 2.5 cm long, the crests to 2 cm. Capsule ca. 3 cm long; seeds flat- tened, + triangular. Chromosome number 2n = 12. Flowering time: (Mid February) March to early July. Distribution: Open woodland or montane grassland in northern Malawi, east- ern Zambia, southwestern Tanzania, and Zaire.—Fic. Moraea macrantha is closely related to M. verdickii and M. textilis (see dis- cussion under M. verdickii). It can be distinguished from M. verdickii by its dark blue flowers, 3-5 bract leaves, and later flowering, from mid February to July; and from M. textilis by its generally larger flowers, with tepals in the 5.7-8 cm range, and anthers 1.2-1.5 cm long. Moraea macrantha is apparently common in northern Malawi and relatively rare elsewhere, though it extends through southwestern Tanzania to the higher areas of Shaba in Zaire. It occurs only in highland areas and flowers from mid- to late-wet season. ALAWI. NORTHERN REGION: Vipya hills, Chapman H 640 (SRGH). Vipya plateau. Salubeni 660 (K, LISC, PRE, SRGH). Vipya link road, Pawek 1060 (K, MAL, SRGH) Vipya, Luwawa, Chapman 1632 (K, SRGH). Mzuzu, Pawek 5396 (K, MO, SRGH), 5832 (MO, SRGH), 2448 (K, MAL). Mzimba district, Jackson 1283 (BM, BR, K, MAL). Rumphi district, Pawek 3595 (K, MAL). Nyika es. Pawek 6696 (MO, PRE). Chitipa, between Misuku hills and Kalenje River, Pawek 5170 (K, MAL, MO, SRGH). 30 km S of Chikangawa, Vipya Mts., Brummitt 10470 (K). Wenya, Benson 1325 (BM). Mafinga Mts. near Chisengo, chan 1864 (SRGH). TANZANIA. SOUTHERN HIGHLANDS: Ulanda, Ulambya, Leedal 599 (EAH). Kasebe, Bundali, Leedal 439 (EAH). ZAÏRE. SHABA: Lukafu, Mutoli- oe Verdick s.n. (BR). Parc Nationale de di nias Pandeluru, de Witte 2611 (BR). Plateau de es e goi Lisowski, Malaisse & Symo 11621 (BR). Kibura Plateau, 25 km N of Mitwaba, Lisowski, Malaisse 4 Symoens 13676 (BR). 5 km S of Poste de Katshupa, dr nA sse d Symoens 5798 (BR). ZAMBIA. EASTERN PROVINCE. Nyika Plateau, Richards 14388 (K). 16. Moraea ventricosa Baker, Bull. Misc. Inform. 1895: 13. 1895. TYPE: Zambia, Northern Province, “Fwambo,” Carson 37/1894 (K, holotype).—Fic. 12B. M. bequaertii De Wild., diea hw? Nov. Regni Veg. 11: 540. 1913. types: Zaire, Shaba (Katanga), Lubumbashi, rt 316 (B, lectotype). Zaire, Tshisenda, Ringoet 419 5 8 syntype); Homblé 9 (BR. 5 Zaire, Lubumbashi, Homblé 238 (BR, syn- oe ). fon medium, 30-45 cm high, unbranched. Corm ca. 1.5 cm in diameter; tunics of pale medium to fine fibers. Prophylls pale to dark brown, broken and becoming fibrous. Leaf exceeding the inflorescence, linear, canaliculate, 3-7 mm wide. Stem bearing 3-5(-6) overlapping, herbaceous bracts. Spathes her- baceous; inner spathe 8-11.5(-13.5) cm long, the outer ca. % the inner. Flowers comparatively small, blue purple or white to pale yellow; outer tepals 4-5(—5.5) cm long, the limb equal or slightly shorter than the claw; inner tepals 3.7-4.5 cm long. Filaments 12-15 mm long, free in the upper third; anthers 8-10(-11) mm long. Ovary ca. 2 em long; style branches 1.4-1.7 cm long, the crests ca. 1 1977] GOLDBLATT—-MORAEA 981 cm long. Capsule ovoid, to 3 cm long; seeds not known. Chromosome number 2 2 Flowering time: Late February to early May (one record from September). Distribution: Burundi, western and southern Tanzania, southeastern Zaire, and northern and western Zambia; open woodland or grassland, often in moister places along the margins of dambos, marshes, etc.—F ic. 13. Moraea ventricosa as treated here includes both white (to pale yellow) and blue, small-flowered plants. It is one of the few species of subgenus Grandiflora where forms with different colored flowers are known. White-flowered plants occur in the eastern part of its range in western Tanzania, Burundi, and north- eastern Zambia, while blue-flowered plants predominate to the west, in Zaire and northwestern Zambia, M. bequaertii from Zaire exemplifying the latter. Though recorded from open woodland or grassland, M. ventricosa is more often found in moist situations as along dambo margins or poorly drained areas. As discussed under M. verdickii, M. ventricosa is part of a complex of closely allied species including M. macrantha and M. textilis. In fact, it appears most closely related to the Angolan M. textilis which generally has larger flowers, with outer tepals in the 5-6 cm range and anthers 10-13.5 mm long. Specimens of M. verdickii have in the past usually been included in M. ventricosa, but the larger yellow flowers of the former, as well as its earlier blooming habit, should be sufficient to prevent confusion. URUNDI. Gitega, Burasira, Lewalle 1728 (BR, K, WAG). Vicinity of Karuzi-Bureru, van der Ben 1966 (K). ANZANIA. SOUTHERN HIGHLANDS: Mtengulu, Kyimbila district, Stolz 2625 (BM, BR, K, PRE). Elton Plateau, S of Chimala, Milton 72 (EAH). Tunduma, Moreau & Moreau 9807 (EA H). Tunduma-Sumbawanga, Leedal 1076 (EAH). WESTERN PROVINCE: Ufipa district, vlei near Zambia border, Whellan 1212 (k, Mir SRGH). Near Chapota, Sum- baw wanga, 5 8507 (K). Lake Kwela, Bullock 2650 (K). Kundi, Bullock s.n. (K). ZA] SHABA: Lubumbashi (Elisabethville), Boitsfort, Robyns 1655 (BR, EAH, WAG). Lubumbashi, Bequaert 316 (BR); Hirschberg 58 (K). Vallée de Lubumbashi, Quarré 3179 (BR, WAG). Kipopo, Thoen 4646 (BR); Schmitz 5835 (BR). Lopoto, Schmitz 4855 (BR). Kafubu, Quarré 182 (BR, WAG). Vallée de Kapiri, Homblé 1238 (BR). Tshinsinka, Manika (Biano) Plateau, Homblé 1275 (BR). AMBIA. NORTHERN PROVINCE; Mbala (Abercorn) district, Lumi River Marsh, . 12278 (SRGH). Kawimbe, Richards 8329 (K), 19038 (K, MO). Kali, dambo, Richar 6 (K). iac PROVINCE: Solwezi, Drummond & Ad Smith 7031 (BR, K, LISC, M, PRE, SRGH). Ndola, Fanshawe 969 (BR, EAH, K, SRGH). Kitwe, Fanshawe 10142 (SRGH); amt 267 (SRGH), 2546 (K). Mutt A River, Lawton 237 (K). Shi- buchinga, Fanshawe 8388 (K, SRGH). Mwinilunga district, Marks 42 (K). Kabompo road, 25 km from Mwinilunga, Edwards 676 ( LISC, PRE, SRGH ). 17. Moraea textilis Baker, Trans. Linn. Soc., London Bot., Ser. 2, 1: 270. 1878. TYPE: Angola, Huila, Lopollo River, flowering in April, Welwitsch 1549 (BM, holotype; COI, K, LISU, isotypes ).—Fic. 12C M. 3 Pax, Bot. Jahrb. Syst. 15: 151. 1893. Type: Angola, Catala Canginga, flower- n Feb., Teuscz in Exped. Mechow 557c (B, 2 BM. isotype ). Puit large, 40-80 cm high, unbranched. Corm ca. 2 cm in diameter; tunics of coarse, grey, wiry fibers. Prophylls rigid, pale, streaked with longitudinal dark veins and irregularly broken. Leaf linear, canaliculate, exceeding the in- 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 | ) — @ VW. ventricosa N ra N q P | . + Lz 4T. h G M. textilis = T ~A | I l | - i dd L | 50 \ - ponen i t F 30 f 0 100 200 300 400 $00 600 mies Í i — n. aa S l | — > 9 200 400 600 800. 1000 KM " — * — —-ę— Ficung 13. Distribution of Moraea ventricosa and M. textilis. florescence, 6-8 mm wide. Stem bearing (3) 57, often dry and usually over- lapping, bract leaves. Spathes herbaceous, becoming dry from above, acute, of- ten becoming reddish; inner spathe 8-15 cm long, the outer ca. * the inner. Flowers blue purple or yellow, usually conspicuously streaked with purple; outer tepals 4.7-6.3 cm long, the claw often slightly exceeding the limb; inner tepals erect, 4.8-6.5 cm long. Filaments 1.5-2.2 cm long, free in the upper quar- 1977] GOLDBLATT—MORAEA 983 ter; anthers 1.0-1.35 cm long. Ovary ca. 2 cm long; style branches 1.5-2.5 cm long, the crests to 1.5 cm long. Capsule ovoid, ca. 3 cm long; seeds flattened and triangular to discoid. Chromosome number not known. Flowering time: (January) April to June (July to August). Distribution: Throughout the Angolan plateau above 1,200 m in grassland or rocky sites, occasionally reported from damper situations.—Fic. 13 A much more restricted concept of M. textilis is proposed here compared to Geerinck's (1970) treatment of this species which included M. ventricosa and M. verdickii. As treated here, M. textilis occurs almost exclusively in Angola, from the highlands in the southwest to the border of Zaire and Zambia, one col- lection being known from western Zambia. Collections suggest that the species is common in the provinces of Huila and Huambo, and relatively rare elsewhere. though this may simply be a reflection of collecting activity being concentrated in population centers. Moraea textilis is reported from grassland and rocky sites at elevations above 1,200 m, though occasionally also from damp places. It has a remarkably long flowering period from December to July, though most col- lections are from the months of May and June, in the dry season. Moraea textilis is most closely related to M. ventricosa, a somewhat smaller species occurring mainly in moist situations to the east, and it is also allied to M. verdickii and M. macrantha as discussed under M. verdickii. The latter in- variably has yellow flowers which are larger than those of M. textilis, and it blooms in the wet season, from November to February and March. BENGUELLA: Highlands between Ganda and Caconda, Hundt 822 (BM, Z). ANG 5 district, homo 1906 (K). Caconda, Gossweiler 4206 (COI). cUANZO SUL: Am- n, Gossweiler er (COI, K). HuAaMno: Huambo (Nova Lisboa), da Silva 3604 (LISC). “sewa dhan Huambo and Quipeio, Exell & Mendonca 1874 (BM). Near Caululo, Moreno 163 (COI, LISC). Cuima, Exell & Mendonça 1937 (BM). Country of the Ganguellas and Am- buellas, Gossweiler 4 (K). Sacaala forest, Murta 170 (COI). Mt. D plateau. above village, Huntley en ) (PRE). una: Hoque, Henriques 1020 (BM, COI, LISC, LISU, SRGH). Leba, Pritchard 339 (BM, K, LISC). Ne = Boa River, beu 1549 (BM, COI, K, LISU). Turdevala NW of Sa de Bandcira, s 3294 (K, LISC, PRE, S); Barbosa & Moreno 10218 (COI). Lubango, sees do Bin E Mandos 3762 (LISC ). Tchivinguiro, Henriques 59 (LISC). LuNpA: R. Coxi, E 91 e Mendonca 1358 (BM). Xassengue Caiango, near River Cuando, Gossweiler 11855 (C OI). w0xico: Texeira de Sousa, Gossweiler 12248, 12249 (BM, LISC) ZAMBIA. WESTERN PROVINCE: Plaius, Mwinilunga district, Marks 42 (K). 18. Moraea tanzanica Goldbl. sp. nov. Type: Tanzania, Southern Highlands, Kyimbila district, Stolz 2142 (K, holotype; BM, BR, GH, PRE, Z, isotypes). —Fic. 14A Planta 20—35 cm alta. Tunicae cormi pallidae. Folium solitarium, canaliculatum, 15-25 on raro spathas excedentium, insertum breviter supra terram. Caulis s ped ferens baat duabus. eo herbaceae, interior 9-10 cm longa, exterior 6-8 cm 11 Flores luteis; 1 exteriora 4.5-5.5 cm longa, limbis 3-3.5 cm longis, expansis; sie 3.5-4.5 m longa. Filamenta 1.2-1.5 cm longa; antherae ca. 1 cm longae. Germen 1.7-2.2 cm longum; rami styli 1.7-2 cm longi; cristae ca. 1.3 cm longae. Plants solitary, medium, 20-35 cm high. Corm ca. 1.5 cm in diameter; tunics of pale medium fibers. Prophylls dry-membranous, pale, irregularly torn above. Leaf solitary, 15-25 cm long, seldom reaching beyond the midline of the spathes, 984 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficung 14. Moraea species—A. M. tanzanica.—B. M. clavata.—C. M. brevifolia. (x0.5). canaliculate, inserted shortly above ground level. Stem simple. Bract leaves 2, usually overlapping. Spathes herbaceous, attenuate; inner spathe 9-10 cm long, the outer ca. % the inner. Flowers yellow; outer tepals 4.5-5.5 cm long, the limb 3-3.5 cm long, spreading; inner tepals erect, 3.5-4.5 cm long. Filaments 1.2-1.5 1977] GOLDBLATT—MORAEA 285 em long, free in the upper half; anthers ca. 1 cm long. Ovary 1.7-2.2 cm long; style branches 1.7-2 cm long, the crests ca. 1.3 cm long. Capsule cylindrical, to 3 em long; seeds flattened, discoid. Chromosome number 2n = 12. Flowering time: January to March. Distribution: Northern Malawi and southern Tanzania, at high altitudes, 2,000-3,000 m; in open mountain grassland.—Fic. 15 Moraea tanzanica is related to the M. textilis complex, and particularly to the yellow-flowered M. verdickii. It is, however, a much smaller species, though with a very large flower, and can always be distinguished from M. verdickii by its consistently short leaf which seldom reaches beyond the midline of the spathes. Moraea tanzanica appears to stand at the beginning of a series of reduced species in which the leaf is generally short and inserted above ground level, sometimes at the base of the inflorescence. In this group the bract leaves are often con- gested and may be reduced in number or even lacking, as in M. unifoliata, the most specialized species in this alliance. The species most closely related to M. tanzanica among the reduced species are M. brevifolia, which often has a shorter leaf and solitary bract leaf, and M. clavata, an early-flowering species with much smaller flowers. Moraea tanzanica is confined to southwestern Tanzania where it has been collected mostly at high altitudes in short grassland. It flowers from January to March, particularly to- wards the end of summer. In contrast, M. brevifolia, found to the west in Zambia, blooms in December to January and occurs only in marshy situations. Moraea clavata, flowering even earlier in late spring to mid summer, is also confined to moist situations and occurs from central Zambia to southern Angola. MALAWI. NORTHERN REGION: Nyika Plateau, Katumbi-Juniper forest turnoff, Pawek 6660 (MO); Phillips 1071A (K, MO, SRGH). TANZANIA. SOUTHERN HIGHLANDS: Kyimbila district, 2142 (BM, BR, GH, K, pui Z). Elton Plateau, Richards 14128 (EAH, K). Rungwe Richards 14314 (K). Rung caldera, Pocs 6505 (DAR). Mbeya Pe sak, Kerfoot 1628 1 Kipengere Mts., Niambie district, Richards 7574, 7617 (K). Ndumbi Forest Reserve, Semsei 1664 (EAH, K). 19. Moraea brevifolia Goldbl., sp. nov. Tyre: Zambia, Northern Region, Lu- mangwe Falls, Mporokoso district, Simon & Williamson 1483 (K, holotype; LISC, SRGH, isotypes ).—Fic. 14C Plantae 25-50 cm alta. Tunicae cormi pallidae, tenues. Folium solitarium, breve, 5-20 cm longum, insertum in pars inferior caulis. Caulis simplex, ferens bractea una, raro dua. Spathae herbaceae, interior 7. 5-12 c m longa, exterior 6-9 cm longa. Flores luteus; tepala ex- teriora 5-7 cm longa, limbis 3-4 cm longis, expansis; interiora 4-5 cm longa. Filamenta 10-12 mm longa; antherae 9-16 mm longae. Germen 1-1.3 cm longum; rami styli 1.2-2.5 cm longi; cristae ad 1.5 cm longas. Plants 25-50 cm high, solitary. Corm 1-1.5 cm in diameter; tunics of fine brown reticulate fibers. Prophylls dry-membranous, becoming lacerated above. Leaf solitary, short, 5-20 cm long, with a free apex or entirely sheathing and bractlike, inserted in the lower half of the stem, above ground level, canaliculate. Stem simple. Bract leaf 1 (occasionally 2), 6-9 cm long. Spathes herbaceous, with brown attenuate apices; inner spathe 7.5-12 cm long, the outer ca. 2-% the inner. Flowers yellow; outer tepals 5-7 cm long, the limb 3-4 cm long, spread- 986 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 / 20 40% aird / / / + M j 30, e % 4 oe ? 20 \ | l| he ND ILLU C73) C i | A T DN 10 Zu 2 . 5 ma UAN \ " ? ° | | L n 10 T | | | | | | | i T | | | | 2 : to @ V. tanzanica | B M. brevifolia L A V. clavata | | | 30 — — | T 0 100200 300 400 $00 600 MILES Ln — 1 , | O 200\400 800 800 1000 KM : — —— \ a w \ | | | ] pe — \ \ \ | | i | | / / L 20 \ 10 Ñ L | | = ie 4 Í s I-10 Ficure 15. Distribution of Moraea tanzanica, M. brevifolia, and M. clavata. ing; inner tepals erect, 4-5 cm long. Filaments 10-12 mm long, free in the up- per third; anthers 9-16 mm long. Ovary 1-1.3 cm long; style branches 1.2-2.5 cm long, the crests to 1.5 cm long. Capsule and seeds unknown. Chromosome number not known. Flowering time: December to January. Distribution: Zambia, central to northeastern Zambia; in wet places.—Fic. 15. 1977] GOLDBLATT—MORAEA 987 Moraea brevifolia is apparently rare and occurs only in marshy situations in northern and western Zambia where it blooms in midsummer. It is somewhat variable in leaf morphology, some plants having a very short, almost entirely sheathing leaf, while in others the leaf is longer with the free portion reaching the spathes. The leaf is, however, distinctive in its point of insertion, well above ground, at about the midline of the stem. Moraea brevifolia is closely related to the predominantly Angolan M. clavata, a generally smaller species, which con- sistently has smaller flowers. Moraea brevifolia may also be confused with M. tanzanica, which has similar flowers, but is vegetatively more robust, has a larger broader leaf, and always has two bract leaves where one is the rule in M. brevi- folia. 'These two species also differ in habitat and time of flowering, M. tanzanica being found in montane grassland and flowering later in the summer than M. brevifolia. ZAMBIA. NORTHERN PROVINCE: Lumangwe Falls, Mporokoso district, Simon & William- son 1483 (K, LISC, SRGH). N'chelengi-Luapula River road, Kawambwa district, Richards 15476 (K). Kambole-Mbala road, Richards 11901 (K). Chinakila- Loye flats, Mbala dis- trict, Richards 19473 (K). WESTERN REGION: Mwinilunga district, Milne- ee 3930 (BM, BR, LISC). Plauis, Mwinilunga district, Marks 41 (K). 20. Moraea clavata Foster, Contr. Gray Herb. 114: 49. 1936. Type: as for M. gracilis Baker.—F Ic. M. gracilis orm hs 7 Linn. Soc. London, Bot., Ser. 2, 1: 271. 1878, hom. illeg. non M. gracilis (Licht. ex Roem. & Schult.) Diels, 1833. Type: Angola, Huila, near Lopollo River, Wed 1545 (BM, lectotype; K, LISU, isolectotypes). Plants small, often growing in clumps, 15-30(-35) cm high. Corm 1-1.5 cm in diameter; tunics of fine, grey brown reticulate fibers. Prophylls short, mem- branous to herbaceous, usually entire at the apices. Leaf solitary, inserted mid- way up the stem, usually 5-12(-20) cm long, rarely exceeding the inflorescence. Stem simple with an extended lower internode. Bract leaf 1, herbaceous, 3-4 cm long, above the leaf. Spathes herbaceous with dry upper margins; inner spathe 4.5-6(-7) cm long, the outer ca. % the inner. Flowers yellow; outer tepals 2-3(-3.5) cm long, the limb 1-2 cm long, spreading; inner tepals 1.3-2.0 cm long, (probably) erect. Filaments 5-6 mm long, united in the lower half; anthers 4-5 mm long. Ovary 5-8 mm long; style branches ca. 1 cm long, the crests to 1.0 em long. Capsule ovoid-globose, 7-10 mm long; seeds small, angular to some- what flattened. Chromosome number not known. Flowering time: October to January ( March). Distribution: Angola and Zambia, in moist situations.—Fic. 15. Moraea clavata was among the first species of Moraea known from tropical Africa, having been collected in the 1850s by Welwitsch during his exploration of Angola. It was first named M. gracilis by Baker (in 1878) but this, a homonym, was replaced with M. clavata by Foster in 1936. Only a few collections are known, mainly from the southwestern highlands of Angola, but there are also records from western and northern Zambia, in the Mwinilunga and Serenje dis- tricts, the former some 800 km and the latter 1,500 km to the east of the Angola populations. It is a moisture loving species, found only in marshy places, dambos, 988 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 etc. and has a very long flowering period, from late spring in October to January, with one record in March. It is a distinctive species, one of the smallest in subgenus Grandiflora, with a rather short leaf inserted well above ground level, a single bract, and a very small flower with an ovary 6-8 mm long. It is clearly related to M. brevifolia, a taller and larger-flowered species which grows in a similar habitat in western and northern Zambia. The difference in size between these two alone makes confusion unlikely. OLA. HUILA: Serra de Chela, Humpata, Gossweiler 13315 (LISC). Humpata Plateau near Nhime, Santos 76 (LISC). Huila, Antunes s.n. (LISU). Near the Lopollo River, Wel- witsch 1545 (BM, COI, K, LISU), Without precise locality, Antunes 643 (L AMBIA. CENTRAL REGION Serenje, Fanshawe 6731 (K, SRGH). Road between Mpika nd Kapiri-Mposhi, Serenje distri ct, Richards 16869 (K). WESTERN REGION: Mwinilunga 1 daabo NE of Dobega Bridge, Milne-Redhead 3620 (K). 21. Moraea upembana Goldbl., sp. nov. Type: Zaïre, Shaba, Parc Nationale de l'Upemba, banks of Lumana River, de Witte 2400 (BR, holotype).— Fic. 16. Planta parva, 10-20 cm alta. Tunicae cormi tenues, reticulatae. Folium ignotum, absens cm longis, expansis; interiora 2-2.5 cm longa. Filamenta ad 7 mm longa; antherae ad 8 mm longas. Germen 8-15 mm longum; rami styli ca. 10 mm longi; cristae ad 7 mm longas. Plants small, 10-20 cm high, unbranched. Corm ca. 1 cm in diameter; tunics of pale to dark, finely reticulate fibers extending upward in a neck. Prophylls dry-membranous, entire or irregularly broken. Leaf unknown, absent at flower- ing time. Stem bearing 1 or 2 herbaceous bract leaves. Spathes herbaceous, with dry, pale brown acute apices; inner spathe 3.5-7 cm long, the outer 3-4 cm long, ca. half the length of the inner. Flowers yellow; outer tepals 2.7-4.5 cm long, the limb 1.5-2.5 cm long, spreading; inner tepals probably erect, 2-2.5 cm long. Filaments to 7 mm long; anthers to 8 mm long. Ovary 8-15 mm long; style branches ca. 10 mm long, the crests to 7 mm long. Capsule and seeds un- known. Chromosome numbers not known. Flowering time: July. Distribution: Zaire, Shaba, apparently very local.—Fic. 17. Known only from the type collection, this rare plant was placed by Geerinck (1970) in his treatment of Moraea in Zaire and Burundi in M. textilis, according to his very wide interpretation of this species. It has however a number of very unusual features which make it appear to me quite distinct and perhaps not even very closely allied to M. textilis. Firstly, M. upembana flowers in July, in the dry season, is relatively short, standing up to 20 cm high, and is apparently quite leafless at this time. In contrast, M. ventricosa and M. macrantha (also included by Geerinck in M. textilis) normally flower in March and April in Shaba at the end of the wet season, and even those plants collected as late as May and June are as tall and as large flowered as the collections from earlier in the season, and do have a well-developed, green leaf. In other morphological features M. upembana also seems distinct, for it has very fine corm tunics, only 1 or 2 short 1977] GOLDBLATT—MORAEA 989 FicurE 16. Type collection of Moraea upembana. bract leaves, and is much smaller in all characteristics than M. ventricosa or M. macrantha, both of which have coarser corm tunics and 3-5 large bract leaves. Though the single collection known is inadequate for a complete understand- ing of M. upembana, it seems best associated with the group of morphologically 990 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 . upembana . bovonei . balundana . unifoliata Distribution of Moraea nene M. bovonei, M. balundana, and M. uni- 1 , FIGURE 17. foliata with morphology of—A. M. bovone M. balundana.—C. M. unifoliata (x0. 5). reduced species of subgenus Grandiflora, and seems most closely related to the Angolan M. clavata and its relative M. brevifolia from Zambia. Further collec- tions are needed to resolve the remaining questions about this species, including its habitat, variation pattern, and even the details of its leaf and flower structure. 1977] GOLDBLATT—MORAEA 291 ZAIRE. SHABA: Parc Nationale de l'Upemba, Lumana River, de Witte 2400 (BR). 22. Moraea bovonei Chiov., Nuovo Giorn. Bot. Ital. 26: 72. 1919. TYPE: Zaire, Shaba, Ditungula, Lake Mweru, Bovone 12 [F, lectotype; T (photograph seen) isolectotype].—F ic. 17A Plants slender, to 40 cm tall, with the leaf extending a further 40 cm. Corm and tunics not known. Prophylls papery, pale, becoming torn and irregular at the apices. Leaf terete, ca. 40 cm long, inserted at the base of the inflorescence. Stem with a long lower internode at the apex of which are 2 herbaceous bract leaves, each ca. 3.5 em long, and the inflorescence. Spathes herbaceous with dry attenuate apices; inner spathe ca. 6 cm long, the outer slightly shorter. Flower yellow; outer tepals to 2.6 cm long; inner tepals to 2.4 cm long. Filaments ‘a. 8 mm long, united in the lower half; anthers 7 mm long. Ovary 8-10 mm long; style branches 1.2 cm long, the crests to 7 mm long. Capsule and seeds un- known. Chromosome number unknown. Flowering time: Summer (February ). Distribution: Zaire, eastern Shaba, marshes of Lake Mweru.—Fic. 17. Moraea bovonei, known from only a single gathering, is a most unusual spe- cies. It has a very long slender stem, some 40 cm high, and the long leaf is in- serted at the stem apex, immediately under the inflorescence. Unlike the re- lated species, M. balundana and M. unifoliata, in which the leaf is also inserted below the inflorescence, the leaf of M. bovonei is very long, approximately equal in length to the stem. Moraea bovonei can be distinguished from its allies not only by its long stem and leaf but also by the presence of two bract leaves at the base of the spathes, whereas there is only one in M. balundana and none in M. unifoliata. ZAIRE. SHABA: Ditungula, Lake Mweru, Bovone 12 ( — ` T). 23. Moraea balundana Goldbl., sp. nov. rype: Zaire, Shaba, 52 km SW of Mutshatsha, Robinson 6025 (K, holotype; M, SRGH, isotypes).—Fic. 17B. Planta 15-40 cm alta. Tunicae cormi fuscae, tenues, reticulatae. Folium solitarium, in- sertum ad base inflorescentiae, 6-11 cm longum, teres. 1 simplex, ferens bracteam unum. luteis; tepala exteriora 3-5 cm longa, limbis 1.7-3 cm longis, expansis; dh interiora 2-3 cm longa. Filamenta 8-10 mm longa; antherae 8-9 mm longae. Germen ca. 8 mm longum; rami styli 1.2-1.5 cm longi; cristae 5-10 mm longae. Plants slender and unbranched, 15-40 cm high. Corm ca. 1.5 cm in diameter; tunics of dark, finely reticulate fibers extending shortly upward in a neck. Prophylls vieil but the upper one dry and becoming fibrous. Leaf 6-11 cm long, terete, inserted at the base of the inflorescence, and shortly exceeding it. Stem slender with a very long lowermost internode. Bract leaf 1, short, brown. Spathes unequal; inner spathe 34.5 cm long, herbaceous below, dry- brown at the apex, the outer 1.5-2 cm long, entirely brown, less than half the inner. Flower yellow; outer tepals 3-5 cm long, the limb 1.7-3 cm long, spread- ing; inner tepals 2-3 cm long, evidently erect. Filaments 8-10 mm long, free in the upper half; anthers 8-9 mm long. Ovary ca. 8 mm long; style branches 992, ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 1.2-1.5 em long, the crests 5-10 mm long. Capsule and seeds unknown. Chromo- some number not known. Flowering time: December. Distribution: Zaire, southwestern Shaba (Katanga) in permanently wet dambo.—F'ic. 17 Moraea balundana, known only from a single gathering in the southern part of Zaire, close to the Angola and Zambia borders, is named after the Balunda tribe who live in this area. The species is closely related to M. bovonei which occurs to the east near Lake Mweru. Moraea balundana differs from M. bovonei mainly in its shorter leaf, single, dry, dark brown bract leaf, and unusually short, brown outer spathe. Further collections are needed to assess the variation in the species, and thus to better understand its relationship to the very similar M. bovonei and to M. unifoliata, also closely related. However, in the absence of more material, the unusual features of leaf, bracts, and spathes necessitate speci- fic recognition. The species was assigned to M. angusta (Thunb.) Ker by Geerinck (1972), but the resemblance to this South African plant from the Cape region is only superficial. There seems no doubt that there is no close relation- ship between the two, assigned by Goldblatt (1976a, 1976b) to different sub- genera of Moraea. ZAIRE. SHABA, 52 km SW of Mutshatsha, near Zambian border, Robinson 6025 (K, M, SRGH ). 24. Moraea unifoliata Foster, Contr. Gray Herb. 114: 48. 1936. Type: as for M. aphylla De Wild.—Fic. 17C. M. aphylla De Wild., Ann. Mus. Congo, Sér. 4, Bot. 2: 21. 1913, hom. illeg. non M. aphylla L.f., 1781. type: Zaire, Shaba (Katanga), Tembwe, Hock s.n. (BR, holotype). Plants 20-35 cm high, unbranched. Corm ca. 1.5 cm wide; tunics of pale medium-fine reticulate fibers extending up in a neck. Prophylls papery, light brown, often fibrous above. Produced leaf of flowering individuals 4-6.5 cm long, seldom longer than the spathes, inserted near the apex of the stem and usually partly sheathing the inflorescence. Stem simple and lacking bract leaves. Spathes 4-6 cm long, herbaceous, or becoming dry, attenuate; inner spathe equal to or shortly exceeding the outer. Flowers yellow; outer tepals lanceolate, ca. 3.5 cm long, the claw to 1.5 cm long, spreading; inner tepals 2.5-3 cm long, erect. Filaments ca. 7.5 mm long, free in the upper part (half to a quarter); anthers 7.5-9 mm long. Ovary 7-10 mm long; style branches to 1.5 cm long, the crests 5-9 mm long. Capsule ovoid-glabose, to 1 cm long, seeds not known. Chromosome number not known. Flowering time: Late November to January. Distribution: Zaire, southern Shaba in moist situations along dambo mar- gins or sponges.—F'c, 17. Moraea unifoliata can immediately be recognized by its apparent lack of a produced leaf. It does, however, have a leaf which is inserted at the base of the inflorescence, but it is very short, and seldom extends more than 6 cm, thus not exceeding the inflorescence spathes. It is also distinctive in lacking bract leaves, 1977] GOLDBLATT—MORAEA 993 a feature which separates it from its close allies, M. bovonei and M. balundana in both of which the leaf is also inserted at the base of the spathes. The slender M. bovonei, however, has a very long leaf and two small bracts, while M. balun- dana has a shorter leaf and a single short, brown bract. These three species all grow in similar situations, moist sponges or along the margins of marshes or dambos, and bloom in summer. Moraea unifoliata is a fairly local species re- corded from several sites, all in southern Shaba near Lumumbashi. The singular habit of M. unifoliata, with its very short produced leaf placed immediately below the inflorescence and its complete lack of bracts, make the species appear one of the most specialized species of subgenus Grandiflora. It appears as the end product of a series of reduced species which begins with M. tanzania and M. brevifolia, and leads with progressive reduction in leaf size and bract leaf number through M. clavata, M. balundana and M. bovonei. AIRE. SHABA (Katanga): Mukuen, Detilleux 175 (BR, WAG). Mukuen Est, Schmitz "m (BR). Matuitui River, 10 km SSE of Lubumbashi (Elisabethville), Schmitz 2371 (BR). Kapeluka, 16 km NE of Lubumbashi, Schmitz 4766 (BR, WAG). Kipopo, 25 km NE of Lubumbashi, Schmitz 8118 (BR, WAG). Marie-José, Quarré 1478 (BR, GH). INCOMPLETELY UNDERSTOOD SPECIES The following taxon, represented by two collections, is problematic and may represent an undescribed species or a variant of a known one. Material is inadequate at present to provide a satisfactory solution. Moraea sp. 1 Two collections only, of a very tall, large yellow-flowered species were made in the region of Lake Tanganyika. One, Robyns 2192 was collected in Zaire, between Pweto and Baudoinville, the other, Wallace 1302, was found in Tanzania. The plants are about 1 m tall and have large flowers which suggests that they may be forms of M. verdickii. They are, however, late flowering, in May, and have between 4 and 7 large bracts which in the Wallace collection are closely imbricate. Typically, M. verdickii blooms from December to Febru- ary and rarely has more than three bracts, which do not overlap. The Robyns collection resembles to some extent M. bella, though it is unusually robust for this species and does not seem to fit in M. bella as presently circumscribed. Further material from this area is needed before a decision can be reached con- cerning these specimens. EXCLUDED SPECIES 1. Moraea revoluta Wright This species is excluded as the type material is teratological. The specimen, said to have been grown at Kew from corms collected in Angola, clearly belongs to subgenus Grandiflora, and the flowers resemble the pale form of M. textilis. Unique for the subgenus, however, are the several, linear, produced leaves. No other species or specimen of subgenus Grandiflora has more than a single leaf and the presence of more is almost certainly abnormal, 994 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 2. Several species of Moraea, described from Angola and Rhodesia, are now recognized as belonging to the genus Ferraria. Some have been transferred to Ferraria while others remain in Moraea. In spite of the multiplicity of names, there is probably only one species of Ferraria in the whole area including Angola, Zambia, Zaire, and Rhodesia. The most commonly accepted name for the species is F. glutinosa (Baker) Rendle and the many synonyms originally placed in Moraea are listed below. Ferraria glutinosa is distinguished from Moraea by several features, the most readily observable being the equitant leaf, while the branched axis with sticky exudate below the internode are features lacking in the tropical African species of Moraea. It is further distinguished from Moraea by its long-lived, tuberlike corms which apparently lack tunics. Ferraria glutinosa (Baker) Rendle, Cat. Afr. Pl. Welw. 2: 27. 1899. Mu 5 Baker, Trans. Linn. Soc. London , Ser. 2, 1: 271. 1878. TYPE: Angola, near Lopollo River, Welwitsch 1543 E 1 aay LISU, isotype). M. e aae Baker, Trans. Linn. Soc. London, Bot., Ser. 2, 1: 271. 1878. TYPE: Angola, Huila, Morro de Lopollo, Welwitsch ii ( BM, “holotype: LISU, oo M. andontenss Baker, Tra inn. Soc. London, Bot., Ser. 2, 1: 271. 1878. PE: Angola, ngo Andongo, Welwitsch 1532 (BM. holotype; LISU, isotype). M. a ee Baker, Trans. Linn. Soc. London Ser Angola, Huila, plains around Humpata and Lopollo, "m 1547 (BM, yes 'LISU, iso- type y M. aurantiaca Baker, Fl. Trop. Afr. 7 (addendum): 575. 1898. rype: Angola, Malange, Mechow 303 ( B, holotype ). M. kitambensis Baker, Fl. Trop. Afr. 5 575. 1898. TYPE: Angola, Bangala, swamps at Kuango, Buchner He (B, holotype M. randii Rendle, J. Bot. 36: 144. 1898. TYPE: “Rhodesia, Bulawayo, Rand s.n. (BM, holo- type Ferraria candelabrum ( Baker) Rendle, Cat. Afr. Pl. MC x 27. 1899. F. andongensis ( Baker) Rendle, Cat. Afr. Pl. Welw. 2: 27. Moraea malangensis Baker, ips Herb. Boissier, sér. z ii 862. 1901. type: Angola, Malange, Mechow 386 (not seen). (Other synonyms include Ferraris pum Baker and F. hirschbergii Bolus.) LITERATURE CITED ins: J. G. 1878. Report on the Liliaceae, Iridaceae, Hypoxidaceae and Haemodoraceae Welwitsch's Angolan herbarium. Trans. Linn. Soc. London, Bot., Ser. 2, 1: 245-274 ————. 1898. gen a Thiselton-Dyer W. T. (editor), Flora of "Tropical Africa. Vol. 7. Reeve & Co., Burtt, B.L. 1 38. 1 carsonii. Bot. Mag. 161: tab. 9544. . B. B. 1974. Karyotype analysis in Moraea and Dietes. Cytologia 39: 525- Cerise D. 1970. Étude du genre Moraea Miller (Iridaceae) au Congo et au Burundi. . Soc. Roy. Bot. Belgique 103: 143-156. —— q 1972. Trois espèces d'Iridacées méconnues au Zaire. Bull. Soc. Roy. Bot. Belgique 5: 5-8. GorLpBLATT, P. 1971. 5 and morphological studies in the southern African Iri- daceae. J. S. African Bot. 37: 317—460. 1973. Contributions to a knowledge of Moraea (lridaceae) in the summer rain- fall region of South Africa. Ann. Missouri Bot. Gard. 60: 204—259. š g hap 5 and subgeneric classification in Moraea (Iridaceae). Ann. Missouri Bot. Gard. 63: 1-23. . 1$ . The genus Moraea in the winter rainfall region of southern Africa. Ann. Missouri Bot. Gard. 63: 657—786. 1977] GOLDBLATT—MORAEA 995 Lewis, W. H. 1966. Chromosomes of two Moraea (Iridaceae) from southern Africa. Sida 1: 381-382. Raven, P. H. 1973. Evolution of Mediterranean floras. Pp. 213-224, in F. di Castri & H. A. Mooney (editors), Mediterranean Type Ecosystems—Origin and Structure. Springer- Verlag, New York. A NEW CLASSIFICATION OF FICUS WILLIAM RAMIREZ B.! ABSTRACT The taxa of Ficus are classified on the basis of the specificity and morphology of their symbiotic wasps ( Agaonidae ), systems of pollination, and morphology and physiology of the figs. The new classification is a modification of i rne iid . with the following changes. In subgenus Ficus, subsection Eriosucea is elevated t ctional rank. Series Rivulares and . do not belong to the group of Nee ee ollinsted figs and are transferred w Cerotosolen-pollinated complex of subgenus Sycomorus. Two subsections, Scabrae an Nonne: are recognized in section Sycidium, and series Phaeopilosae and subsection Paleomorphe are recognized as sections. The subgenus Sycomorus is much expanded to in- clude eight sections: dat Neomorphe, Prostratae, Pungentes, Pseudopalmae, Rivu- lares, Sycocarpus, and Sycomoru The object of this study is to group the taxa of Ficus into related groups considering the specificity and morphology of their symbiotic agaonids, the dif- ferent systems of pollination, as well as the morphology and physiology of the figs. The last systematic arrangement of Ficus was made by Corner (1965) and is summarized in Table 1. A parallel list of the pollinating agaonids (genera or subgenera reported up to now for each fig taxon) is also included. The list of agaonids was taken from Hill (1967) and modified by me. Parallel to the groups of wasps there are columns showing the absence or presence of corbiculae in the wasps ( Ramirez, 1974). THe NEW CLASSIFICATION OF FICUS AND ITs POLLINATORS The proposed classification of Ficus is found in Table 3. Modifications are extended only to the level of series. SUBGENUS UROSTIGMA This group of figs remains as treated by Corner (1965) (Table 1). Section Urostigma.—The figs are inhabited by Blastophaga (group E), which are characterized by the presence of coxal and sternal corbiculae (as in Figs. 3 and 4). Section Leucogyne.—This section comprises two species. One of them (F tsiela) is pollinated by Maniella delhiensis, with coxal and sternal corbiculae (as in Figs. 3 and 4). Section Conosycea.—The species of this section are pollinated by several groups of wasps. The only Blastophaga (B. arnottiana and B. errata) known from this group of figs have sternal corbiculae and coxal combs. Ceratosolen megarho- palus (the Megarhopalus group) and the majority of Waterstoniella wasps are characterized by only very rudimentary sternal corbiculae (Figs. 5-6); some ! Facultad de Agronomía, Universidad de Costa Rica, Costa Rica, América Central. ANN. Missouni Bor. Garp. 64: 296-310. 1977. 1977] RAMIREZ CLASSIFICATION OF FICUS 997 Waterstoniella (e.g., W. sundaica and W. jacobsoni) do not have corbiculae (Fig. 2). The other two groups of agaonids (Eupristina and Parapristina) found in section Conosycea have sternal and coxal corbiculae (as in Figs. 3 and Section Stilpnophyllum.—This section contains only Ficus elastica which is pollinated by Blastophaga clavigera ( Blastophaga group B) a wasp with sternal and presumably coxal corbiculae (Wiebes, personal communication). Section Malvanthera.—This group is unique in that its anthers have two pol- len sacs which dehisce with one crescentic or equatorial slit. The section is pol- linated by Pleistodontes wasps. However, there are apparently several Pleis- todontes groups pollinating the different groups of Malvanthera figs (personal observation ). Pleistodontes imperialis is characterized by sternal and possibly coxal cor- biculae while the other known Pleistodontes do not possess corbiculae at all. Series Malvanthereae is pollinated by wasps without corbiculae ( P. blandus, frog- gatti, rieki, plebejus, and regalis). The only Pleistodontes (P. inmaturus) known from series Cyclanthereae apparently does not possess corbiculae. For more in- formation on Pleistodontes wasps see Wiebes (1963b: 319, Table 1). It is probable that the group Pleistodontes as well as its Ficus hosts, will have to be reclassified when more is known of both groups. Section Galoglychia.—This group of figs resembles the last section in the in- flexed, not interlocking, apical and internal bracts of the ostiolum (Corner 1959: 376), but it has normal anthers with four pollen sacs. It is pollinated by two main groups of wasps: (a) those with only sternal corbiculae ( Agaon, Allotrio- zoon and Paragaon) and (b) those with sternal and coxal corbiculae ( Alfon- siella and Elisabethiella ). Section Americana.—According to Corner (1959: 376) this section is closely related to both sections Urostigma and Conosycea. It is pollinated by Blastophaga wasps of the subgenus Pegoscapus (Ramirez, 1970) with coxal and sternal corbiculae. However, P. carlosi and P. mariae (the pollinators of F. tuerckheimii in Costa Rica, Mexico and Panama) do not possess coxal corbiculae ( Ramírez, 1970). SUBGENUS PHARMACOSYCEA Corner (1959: 407) considered that the Old World section Oreosycea has the same essential characters, and indeed, is with difficulty distinguished from New World Pharmacosycea species. However, in the descriptions of the two sec- tions we find very important differences, some of which are pointed out in Table 2 The Old World species have in the past been referred to the subgenus Uro- stigma where they are out of place, particularly in being independent trees and not banyans or stranglers. The species from New Caledonia have never been properly classified and they are the closest in several respects to the American species. Corner (1959: 407) stated that he divides the subgenus Pharmacosycea into two sections, maintaining the geographical distinction for convenience, but that redefinition will be necessary when the American species are better known. [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN 298 pupnotiƏtuy + + sndpososadg + + pyarsuofiy + + »]Jonpiaqsi — uoozot4qo]]V — uop ND t — uopdy D12fi]80]pz) + $21uopojsi2]d 2D2421]uD]9fi?) + saquopojsia}d sauuofnpiuadsog "ES + səluopo1s12]d apapodfijn]d + $21uopojsia]d 202421JuDajpq + Sə1uopols1ə]d apywazIDAqny Əpə4ƏtJ1upq]p W p4Əu1upq]p W d + papydoj sn) g wunyâydoudms 4 J nuisiudpand ana]pfido]]p?) + + nuijsiudn] apnaunupluag punupluəaç] 2p1niofiədq + 7]]21u01$4230 M appypaqns q 2Duu442qv]25) uoinauohja1q + D]]Ə1uO1S191D M — 7]]21u03s4121D M + p8mjdolsp]gq + — puysudny Əpəuupuatsspai D) + D]]21u03s423D M 2D91211308fi7 + + puqqsudn:] ƏpƏ1pu] + + Duijsiudnz] 2naopdna(] anaopdna(q + dnoi8 sn[edoure29]q d ＋ + papydoj}sy] g 2Dpi]pA pəoñsouo> pəoñsouop + + pauo W aufigoonor] d + papydojsy] gq Əp.nəuot]11 O + — g pijd os əpñuqoqol]npn5 + + pavydo}sn] g apqsadng — + p5mjdolsn]g 2DSO121]24 DUWS14SO1 f] DUBUSOLN ]pXo) pwəag qussqyv 9epruoes y souosqug sonog uomnoesqng uo snuasqns ov[notq107) '(G96T ) 19uioO uro1j UIJL} sr SNAHA jo u«op»ea1q oneurajs4s oup (2961 “IH Wo1} pegrpour) spruoese jo sdnois juo19jjgrp Suoure ov[noIq100 [eul9js pur [exo jo souasqe 10 9ouoso1d ay} 0} 92u219j81 1e[non1ed u ‘sno jo sdnois ju919jjrp 0j (vQ1ouosqns pue *19uo8) ovpruoegy jo Ayorpioeds jsou poepioosuq 'I ATAV], 299 LASSIFICATION OF FICUS ` RAMÍREZ-—C 1977] IDYIYSIG] anopiouonsiqq aDSOIUDLY IDIDADIXT 2D22D]u2tDM $2]upjua] apnjounjpg apupluojung IDIIDJUIWDY apaydosjouos0g + papydoj sp] gq 20219DUIB1S012D]d 2D0291JDWU1J8018D]d snppjoozn[y aDAAPUDUO TY IDIDANY 2pn1piny apyofiauny SIJUIDISI YI ADIDSOY IIL J, + DBpwudojspig 9poolisoi"] anoofisou'^] n2ofisotg appdogspg + pop dolsp]ç apoofisopod anaofisopoqd + p5ondolsp]gq Əpəuñpo.n1ñag + Dnapudojspig 2D2321D7) 3pDaofisoutg + U310S$010942 J Əpəuu]pdopnəsq SƏə4p]nan] Snan Snag 81024 + sndD4Ja I, Daofisoopuuntu[d + + popdolspn]q ƏpoƏ1uopə]poo.1]sny + + p5pt|dolsp]gq apsoasa N + + popydojsn)g + + Suymoq Əpso]nƏspA D32[i$02 anpndiqy 2D$0]Dn2SD A D22fis$024() -DULIDY J Įex09 peus quosqy əLpiuors y sərrəsq"nç SƏLI9S uolpasqng uonoog snuasqns əuv|norqio5 (ponuquoq) I Av L [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN 300 + u23]08s03D127) aofispndpng Sid oi. Sasuat A + u2]0s03D127) sapiojspaudoony L sapio4pufi?) pnuiadsinouny snda4n2o09fig IDIDIUIID'T + uə]oso1pi9D 2D]DB2MDA SNA + uajosowsa J 2Dgjpjnoiuny — uə]oso1niəD aydsowoa N + u2]0803D427) hug od — u2]0$03D427) apuasiyduy pusadsouapy ES wunpodoyssodrT əm]ofisoiq14 + unpodoyssodrv] 2D40]fingnu1jy — wunjodoyssodrT anjppidsn;) + unpodoyssodry] Əp1n]nqns + unjodoyssodry] pv d, Ət]Jduouuo2]Dd — napnudojsp]g 2DjD12dspx'] avayfiydophg avapfiydosaja H + hi + pavydojsv]g IDIQDIS + d pijdosspiꝗ apsoido;) + ,Danydoj4sn] g ,avsopdoapyd + u23]0$03D427) saquasung + u3]0503D427) ODIDA4SOLd uunipiofis tumpiñs 8014 uunipiofisoutg Imaan Iny apyojaung apyoyoung anadapooidy + ponjdolsn]g 29fiso]DN anadapn201214 |. Iro) [euis qussqv oPpruoez y souosqng səuiəç uonoesqnug uonoog snuogqug 9?[norq107) (penunuo)) I x18v[ 301 RAMÍREZ—CLASSIFICATION OF FICUS 1977] `( LOGT ‘IMH) Ssə[euuəj ay} ur 4XI[[eroədsə 'uojosojpi2;) yya Surneprisiojur Jo səərñəp ſuuν,lñ np moys I} ynq 'papudolsmpng ut pooe[d aie sawads ƏSƏ} luəsəid 1V x + u23]0$03D427) snsowoohg 2D]D|no19sD[ no qu. apaid.ip90ar) + 1008070 apjynplayn 4 + 90/0807 sə4n]l1xyv + U310$04D42 O əppidsiH — u2310$0)D42;) 2Djs28u0/7) + u3J0$0D427) anuimdo]p?) SOJUDISADL d avp nasv] -nouaqn p anjpjnouaqnj3asuo'r] sndap202fig D]fi]js049D JA] snjopida'] [PXo puas jussqy oepruoea y sauasqug Sog uonoasqug 01098 snussqus əv[norqioO (penunuo)) I alavy 302 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 — ma * J 1 A gs 5 Ficur dope ns without corbiculae of „ DEN the urea of the Sdible fig.— 'sosternum of Blastophaga ( Waterstoniella) sundaica, a wasp withot corbiculae but Sd r) bristles which are ; used to carr 1 —3. Front ie of Maniella delhiensis with coxal corbicula.—4 side of mesosternum of Blastophaga (Pegoscapus) cumanensis showing corbicula id some ala in place Mesosternum of sa (Waterstoniella) sundaica with in corbiculae.—6. Mes 'osternum of Cera- tosolen megarhopalus (the Megarhopalus group), right corbicula with some pollen.—7. Meso- sternum of Blastophaga jacana PM A group B) with developed open corbiculae, —8. Mesosternum of Liporrhopalum mindanaensis with closed corbiculae as in Ceratoso 1977] RAMÍREZ— CLASSIFICATION OF FICUS 303 TABLE 2. Presence or absence of some characters in the two sections of subgenus Pharmacosycea. “Pollen Figs Ostiolum Ovary ie Pachycaulous Anther Ex Section Single Crate riform a Re d Mar rees Numerous at t Male Phase Phaku osycea + + + = + m Oreosycea _ = = 4 T — Corner (1967: 40) noted that the new look brought into the subgenus Pharmacosycea by the plants from New Caledonia is the brown hairiness, some- times almost furriness, of twig, leaf, and fig, coupled with the rosettes of large leaves, the many-veined obovate lamina with cordate base and short petiole, and the large size of the fig. All of these characters are more or less primitive and pachycaul signs in Ficus. Section Pharmacosycea in the New World does not present all the pachycaulous characters mentioned by Corner (1967) for some Old World Oreosycea. In order to explain the presence of pharmacosyceous figs in both the Old and New World, Corner (1967: 41) postulated that there must have been a land connection with tropical Africa such as is suggested by the great extension of the 4,000 mile line to the west of Peru. In 1967 he further stated that this con- nection is demanded by other moraceous genera such as Antiaris, Antiaropsis, Sparattosyce and Trophis, as well as by the monocotyledons Dianella, Heliconia and Spathiphyllum in very diverse families. Two hypotheses to explain the presence of Pharmacosycea in the Old and New Worlds are: (a) Sections Pharmacosycea and Oreosycea do not belong to the same subgeneric taxon and their species are more or less similar because of convergence. If this is true, each should be elevated to the subgeneric level, forming biological units separated geographically and by their respective pol- linators, New World Pharmacosycea being the host of Tetrapus wasps (without corbiculae) and Old World Oreosycea of Blastophaga (Blastophaga group F) and of Dolichoris vasculosae (both with coxal and sternal corbiculae). (b) Sec- tions Pharmacosycea and Oreosycea belong to the same subgeneric category, but section Pharmacosycea migrated to the New World before the agaonids evolved corbiculae. This line of thought would agree with the ideas of Corner (1967: 53), although not demonstrating the particular land connection that he postulated. SUBGENUS FICUS In the new classification the subsections Ficus and Eriosycea are elevated to sectional rank as suggested by Corner (1959: 417). The series Rivulares and Pseudopalmeae are not considered to belong to the group of Blastophaga-pol- linated figs and are transferred to the new Ceratosolen-pollinated complex (the subgenus Sycomorus, Table 3). Corner (1969b: 326) stated that F. pseudopalma and F. rivularis (two Philippine species) differ from the rest of section Ficus and from each other markedly enough to require separate taxonomic series (Table 304 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 E 3. Proposed n of the genus Ficus considering the specificity and N of its symbiotic agaonids, the 5 systems of pollination, as well as the morphology and E WA ines of the figs; with a list of the agaonid pollinators (modified from Hill, 1967) of each group, and the presence or absence of corbiculae Corbiculae Subgenus Section Subsection Agi aonidae Absent Sternal Coxal Urostigma aden Blastophaga + + Group E Leucogyne 979 + + Conosycea Conosycea ** + +? si elon ana + Group Eupristina T * Waterstoniella Waterstoniella T Dictyoneuron Waterstoniella + Eupristina + Benjamina Parapristina + + Stilpnophyllum Blastophaga +? clavigera (=Blastophaga Group G) Malvanthera Pleistodontes + Pleistodontes + +? Galoclychia Agaon + Alfonsiella + Allotriozoon + Elisabethiella + + Paragaon + Americana Pegoscapus + + Pharma- Oreosycea Blastophaga + + cosycea y Dolichoris + + Pharmacosycea Tetrapus + Ficus Ficus Blastophaga + sroup Rhizocladus Blastophaga + sroup A Kalosyce Blastophaga 4 Group A Pic ad Erio Blastophaga + B Sycidium Scabrae Blastophaga + Group B Varinga Blastophaga + roup Phaeopilosae Blastophaga + roup Paleomorphe Paleomorphe Liporrhopalum 4 + Copiosae Blastophaga Group D 1977] RAMÍREZ—CLASSIFICATION OF FICUS 305 TABLE 3. (Continued ) Corbiculae Subgenus Section Subsection Agaonidae Absent Sternal Coxal Sucomorus Adenosperma Ceratosolen + omorphe Ceratosolen + Prostratae Ceratosolen + Pugentes Ceratosolen + 6 Ceratosolen + Rivula + d Ceratosolen + Sycocarpus Ceratosolen + a Probably pollinated by a wasp of in ea Group A. > Probably pollinated by a Ceratosolen wa 1). Wiebes (1963a: 101, 104) indicated that the pollinator of F. pseudopalma (C. bakeri) has aberrant characters for the genus Ceratosolen, but appears re- lated to the C. abnormis and C. armipes groups (pollinators of figs of section Sycocarpus ). Sections Kalosyce and Rhizocladus.—These two sections are left in the taxo- nomic position given them by Corner (1965). They form two well-defined groups pollinated by Blastophaga (Blastophaga group A) wasps without corbiculae (Fig. 1). The pollinators of these two groups of figs are quite similar to the ones found with section Ficus (Table 3). These two sections are associated by their pollinators. Corner (1960: 3), however, suggested that sections Kalosyce and Rhizocladus might be considered to form a fifth subgenus. Section Sinosycidium.—This section is left in the same taxonomic position given by Corner (1960: 24). It has a single species (F. tsiangii). Because of its dispersed diandrous flowers and the slightly bifid stigmata of the female flowers, I consider this section to be related to section Ficus (as in Table 3), although the ramiflorous bracteate receptables are like those which occur in sections Sycidium, Sycocarpus and Adenosperma according to Corner (1960: 24-25). The pollinator of F. tsiangii is not known, but it could be a Blastophaga without corbiculae (as in Fig. 1) as those of Blastophaga group A. Section Sycidium.—In the new classification this group has two subsections, Scabrae and Varinga. These groups are related by their pollinators of the Blastophaga group B, which are characterized by their open sternal corbiculae (Fig. 7 Sections Phaeopilosae and Paleomorphe.—The series Phaeopilosae and sub- section Paleomorphe (both sensu Corner, 1965) are elevated to sectional rank. Phaeopilosae is pollinated by Blastophaga group C with closed sternal corbic- ulae (Fig. 9). Paleomorphe has two subsections, Paleomorphe being pollinated by Liporrhopalum with closed sternal corbiculae (Fig. 8) and Copiosae (series Copiosae, sensu Corner, 1965) by Blastophaga group D having closed sternal corbiculae (as in Fig. 9). 306 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 SUBGENUS SYCOMORUS In the new classification the subgenus Sycomorus is expanded and comprises eight sections: Adenosperma, Neomorphe, Prostratae, Pungentes, Pseudopal- meae, Rivulares, Sycocarpus and Sycomorus (Table 3). Of these sections, Adenosperma, Neomorphe and Sycocarpus were considered by Corner (1965) as sections of the subgenus Ficus; Prostratae and Pungentes as series of subsec- tion Sycidium; Pseudopalmeae and Rivulares as series of subsection Ficus. All the sections included here in Sycomorus, excepting Rivulares, are known to be pollinated by Ceratosolen wasps. The pollinator of Ficus rivularis (the only species of section Rivulares) is not known, but I suspect this species to be pollinated by a Ceratosolen with a short ovipositor and closed sternal corbiculae. All the dioecious sections (Adenosperma, Neomorphe, Prostratae, Pungentes, Pseudopalmeae and Sycocarpus) are inhabited by Ceratosolen wasps with short ovipositors. Nevertheless, Corner (1965: 85) included in section Sycocarpus (subsection Papuasyce) the species F. microdictya (of New Guinea) which has the perianth similar to that of Sycocarpus, but is monoecious like Sycomorus?, which does not occur in New Guinea (Corner, 1958: 31, personal communication). Section Sycomorus is a monoecious group pollinated by Ceratosolen with long ovi- positors. RELATIONSHIPS AMONG Groups OF FIGs INCLUDED IN SUBGENUS SYCOMORUS SECTION ADENOSPERMA This section aligns with the unistaminate sections Sycidium and Sycocarpus, which differ in the form of the seed if not in that of the flower (Corner, 1969b: 320). The section is related to section Sycocarpus, subsection Auriculisperma, of the Solomon Islands, and connects with the origin of section Ficus r the Philippine species F. pseudopalma and F. rivularis (Corner, 1969b: 319). SECTION NEOMORPHE Corner (1967: 51) stated that this section has much in common with the sub- genus Sycomorus. Neomorphe may have come from the stock of Adenosperma on the Melanesian Foreland, and this stock may have been connected with that of Sycomorus, so that Neomorphe is an eastern parallel of it (Corner, 1967: 51). Neomorphe must be divided into two series (Table 1), Variegatae and Auricula- tae, which show alliance with the subgenus Sycomorus in the first case and sec- tion Sycocarpus in the second. Series Variegatae can be divided, like- wise, into two subseries. The first Corner (1965: 32-33) called subseries Laciniatae. It has tepals characteristic of subgenus Sycomorus, but it is fur- ther removed geographically from the African subgenus Sycomorus (Cor- ner, 1967). “The second, subseries Variegatae, has only two spe- cies, F. variegata and F. viridicarpa. Ceratosolen striatus (=C. appendiculatus), an agaonid collected from F. variegata in Java, was illustrated by Grandi (1917: ? Ficus pritchardii, a monoecious fig, also belongs to Sycocarpus ( Corner, 1970). 1977] RAMÍREZ—CLASSIFICATION OF FICUS 307 Fig. XII, 6) as a wasp with a long ovipositor like the wasps found in section Sycomorus (as in Table 3). Neomorphe as well as subgenus Sycomorus of Corner (1965) are pollinated by Ceratosolen wasps which are apparently related. Wiebes (1963a: 104) re- ported that the species of the Ceratosolen appendiculatus group live in the re- ceptables of section Neomorphe and subgenus Sycomorus (sensu Corner, 1965), and one species is known from series Prostratae. The occurrence of a group of such closely related species of Ceratosolen in the figs of both dioecious Neomorphe and monoecious Sycomorus would suggest that the floral characters in which Neomorphe is close to Sycomorus are more important than the distribution of the flowers in the receptacles. A parallel is found in F. microdictya, which is a mon- oecious species in the dioecious Sycocarpus? (Wiebes, 1963a: 104 SECTIONS PROSTRATAE AND PUNGENTES These sections are also pollinated by Ceratosolen wasps. Corner (1965) con- siders them to be two series of section Sycidium. According to Wiebes (1963a: 102), the greater part of the Indomalayan and Papuan species of Ceratosolen live in the sections Neomorphe and Sycocarpus, but some are known from Prostratae and Pungentes, two series of Sycidium (sensu Corner, 1965). These series have usually been placed in section Sycocarpus and only recently have been assigned to Sycidium (Wiebes, 1963a). Botanically these two series point to a common ancestor which would combine Sycidium with Sycocarpus and Sycomorus, in- cluding Neomorphe (Corner, 1958: 31). In the opinion of Wiebes (1963a: 102) the wasps from the series Prostratae connect those from the section Neomorphe with those of the subgenus Sycomorus, and the wasps from the series Pungentes appear to be related to the wasps from the section Sycocarpus. According to Corner (1959: 444), series Prostratae relates with section Ficus but habit and convenience place it in Sycidium. SECTIONS PSEUDOPALMEAE AND RIVULARES Each of these taxa has a single species. Corner (1965) included them as two series of the subgenus Ficus. Both species are found in the Philippines. Ficus rivularis is an advanced leptocaul shrub with lanceolate leaves, distinguished in section Ficus by the gamophyllous perianth with distinct tepal lobes, com- pressed auriculiform seed, and the more or less gynobasic style in the female flower. The perianth is intermediate between that of section Ficus and Syco- carpus. In perianth, style and seed, F. rivularis agrees with Adenosperma; it ap- pears as a relic, fitting no section of the ancestral line of section Ficus from which those of Auriculisperma and Adenosperma diverged (Corner 1969b: 328). Ficus pseudopalma connects as a pachycaul with F. dammaropsis (section Sycocarpus, subsection Auriculisperma) of New Guinea, and thus, with section Adenosperma. It connects also with the ancestry of the F. deltoidea complex (section Ficus series Erythrogyneae) and has the three tepals of section Ficus * Ficus pritchardii (a monoecious fig) also belongs to Sycocarpus. 308 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 FicunEs 9-10.—9. Mesosternum of Blastophaga jacobsi (Blastophaga group C) with closed sternal corbiculae.—10. Mesosternum of Ceratosolen pilipis with closed corbiculae. ( Corner, 1969b: 326). Ceratosolen bakeri is the pollinator of F. pseudopalma. This wasp appears to be related to the C. abnormis and the C. armipes groups. Ficus pseudopalma was classified in section Ficus because of its bistaminate male flowers, but it does show some relationship with F. dammaropsis (section Sycocarpus), the host of C. abnormis (Wiebes, 1963a: 101). SECTION SYCOCARPUS This group of Ficus is mostly dioecious; however, F. microdictya and F. pritchardii are monoecious. It is pollinated by Ceratosolen wasps with short ovipositors, but the ovipositors of the pollinators of F. microdictya and F. pritchar- dii are probably much longer than the abdomens. The chief character of the section is the entirely gamophyllous perianth. In the male flower the perianth is saccate and covers one, or less often, two stamens (Corner 1960: 38). For the relationship of the pollinators of Sycocarpus with the pollinator of F. pseudo- palma and those of section Nemorphe, see under sections Pseudopalmeae and Neomorphe. See also under section Adenosperma. SECTION SYCOMORUS In the new classification, this group contains all the monoecious figs in- cluded in the subgenus Sycomorus of Corner (1965). It is pollinated by Cerato- solen wasps with long ovipositors. Galil (1973) noted that in spite of numerous structural differences between the syconia of the dioecious F. fistulosa (section Sycocarpus) and the monoe- cious F. sycomorus (section Sycomorus sensu Ramírez, 1974) which belong to different subgenera of Ficus, namely Ficus and Sycomorus (sensu Corner, 1965) respectively, the two have several biological features in common. In both, the pollinating wasps are species of Ceratosolen which behave very similarly in re- lation to the figs, and such likeness in behavior indicates that physiological con- ditions within the figs are probably also similar in both cases. 1977] RAMIREZ CLASSIFICATION OF FICUS 309 CHARACTERS OF THE SUBGENUS SYCOMORUS Corner (1967: 51) stated that Sycomorus, Sycocarpus, Adenosperma, Neo- morphe, and two series of Sycidium (Prostratae and Pungentes) are distinguished by having Ceratosolen as pollinating insects. Despite their differences, he sug- gests it may be necessary to combine them in the subgenus Sycomorus in con- trast with the remainder of the subgenus Ficus pollinated by Blastophaga. The newly defined subgenus Sycomorus is characterized by the following characters: Male flowers: (a) in 1 or 2 (in some cases 3) ostiolar rings; (b) few per fig; (c) usually without pistillode; (d) perianth with free petals, gamo- phyllous or utriculate; (e) mostly sessile; (f) usually with only one or two stamens (few species with three). Anthers: (a) enfolded by the perianth; (b) usually small; (c) pollen not exposed at male phase. Female flowers: (a) stigma simple; (b) styles usually short excepting those of section Sycomorus and of F. microdictya and pritchardii*. Syconia: (a) with internal bristles; (b) helicoidal ostiolar entrance with several (more than three) interleafing super- ficial bracts; (c) dioecious (excepting section Sycomorus and F. microdictya and pritchardii; (d) ostiolum usually does not open at male phase. Leaf: (a stomata usually superficial; (b) leaf not coriaceous; (c) plicate in bud. Trees: independent, not epiphytic. Pollinators: Ceratosolen wasps which are character- ized by closed sternal corbiculae (as in Fig. 10), and coxal combs, and which col- lect the pollen from detached anthers cut by the males (Galil, 1973); short ovipositors (except the Ceratosolen wasps of section Sycomorus and F. micro- dictya and pritchardii) and by the ability of the male to perforate the fig in order to gnaw an exit that allows the females to escape. The males in all species probably cut the stamens before the females emerge from the galls (Galil, 1973). The figs of sections Adenosperma, Sycocarpus, and Sycomorus are para- sitized by Eukoebelea wasps (tribe Sycophagini, Hill, 1967: 92), while the spe- cies of. section Sycomorus are inhabited by Sycophaga wasps (tribe Sycophagini, Hill, 1967: 92). LITERATURE CITED Corner, E. T H. 1958. An neu to the distribution of Ficus. Reinwardtia 4: 15-45. Taxonomic notes on Ficus Linn., Asia and Australasia. Sections 1-4. Gard. Bull. Singapore 17: 368-485. 1960. Taxonomic notes on Ficus Linn., Asia and Australasia. Sections 5-6. Gard. Bull. SIME 18: 1—69. 1965. Check-list of Ficus in Asia and Australasia with keys to identification. Gard. Bull. recen 21: 1-1 967. Ficus in the Solomon 2 p its bearing on the Post-Jurassic History of Melanesia. Philos. Trans B, a. The p ‘of rod 9e a recent invasion of the Sunda Shelf. Philos. Trans., Ser. B, 256: 281-317. 19 Ficus section Adenosperma. Philos. Trans., Ser. B, 256: 319—355. E Ficus subgen. Pharmacosycea with 1 to the species of New Cale- donia. Philos. Trans., Ser. B, 259: 383.433. 3 . 383) suggested that subsection . (of section Sycocarpus), to which F. na, microdictya and pritchardii belong, should become a fifth subgenus as a monoecious 1 distinct from subgenus Ficus but with F. itoana as the dioecious product. 310 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 GALIL, J. 1973. Pollination in BSa figs; pollination of Ficus fistulosa by Ceratosolen hewitti. Gard. Bull. Singapore 26: 303-311. GRANDI, G. 1917. ae "alla conoscenza degli Agaonini ( Hymenoptera, Chalcididae ) i Giava. Boll. Lab. Zool. Gen. Agrar. R. Scuola Agric. Portici 12: 1-60. Hix, E = 1967. Figs (Ficus spp.) of Hong Kong. Hong Kong University Press, Hong 128 pp. Pub B., W. 1970. Host specificity of fig wasps ( Agaonidae). Evolution 24: 680-691. 9 4. Coevolution of Ficus and Agaonidae. Ann. Missouri Bot. Garden 61: 770- WIEBEs, LT 963a. Taxonomy and host preferences of n Australian fig wasps of the genus Ceratosolen (Agaonidae). Tijdschr. Entomol. 106: 1-112. ——. 1963b. Indo-Malayan and Papuan fig wasps 1 aoe. epee 2. The genus Pleistodontes Saunders 1 Zool. Meded., Leiden 38: 303-321. STUDIES IN BIGNONIACEAE 25: NEW SPECIES AND COMBINATIONS IN SOUTH AMERICAN BIGNONIACEAE' ALWYN H. GENTRY" ABSTRACT Five new species of South American Bignoniaceae are descri 5 gran- villei A. Gentry, Arrabidaea ornithophila A. Gentry, Cuspidaria octoptera A. Gentry, Memora cristicalyx A. Gent and Tabebuia catarinensis A. Gentry—and two new co 1 Lundia virginalis var. "nitidula (DC.) A. Gentry and Memora imperatoris-maximilianii ( Wawra ) . Gentry—are made. Anemopaegma granvillei A. Gentry, sp nov. utex scandens; sine pseudostipulis vel consociebus gl e in nodis inter petiolos; folia 1 foliolis oblongo-ellipticis, infra omnino puberulis; inflorescentia axillaris, ra sa, puberula; flores iue 1 corolla lutea, tubo pios Eo ovario ovoideo, cei ah lepidoto, ad basim non contracto; fructus ignotus. Vine; branchlets finely but prominently striate, elenticellate, puberulous, without interpetiolar glandular fields; pseudostipules (only 1 seen) subulate, 4 mm by 1 mm. Leaves 2-foliolate, sometimes tendrillate (tendril tip not seen); leaflets (ovate-)oblong-elliptic, shortly and obtusely acuminate, rounded or truncate at the base, 10-15 cm long, 4.5-6.5 cm wide, chartaceous, puberulous throughout beneath with rather scattered erect trichomes, above puberulous only along the main veins, the secondary veins looped and connecting several mm from the margins, not very prominent nor strongly ascending, drying olive, glossy above, dull below. Inflorescence a contracted axillary raceme, densely tannish puberulous; pedicels subtended by linear 2-3 mm long bracts. Flowers with the calyx cupular, asymmetrically truncate, 7-10 mm long, 7-8 mm wide, puberulous at the base and around the margin, with fields of plate-shaped glands in the upper half; corolla tube pale to lemon yellow, the lobes pale yellow, tubular-campanulate, ca. 5 cm long, the tube 3.5-4 cm long, the lobes ca. 1 cm long, the tube glabrous outside, the lobes glandular-lepidote with ciliate mar- gins, large glands absent below the lobes; stamens didynamous; ovary (in bud) ovoid, longitudinally ridged, minutely lepidote, not contracted at the base; disc large, patelliform, 2 mm long, 3 mm wide. Fruit unknown. Type: FRENCH GuIANA. Riviere Petite Ouaqui, végétation ripicole verse l'embouchure de la crique Carbet Brülé, 27 July 1973, de Granville 1935 (CAY, holotype and isotype; MO, fragments and photocopy ). The combination of glabrous corolla tube and puberulous leaves indicates affinity with A. puberulum (Seib.) Miranda which belongs to the A. grandi- Tuin. (aed Merrill & Sandw. complex. However, all species of this complex Supported by grants from the National Science Foundation. ? Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. ANN. Missouni Bor. Garp. 64: 311-319. 1977. 312 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 differ in having more prominently lenticellate branchlets and much more promi- nent and strongly ascending secondary venation. Inflorescence bracts of A. puberulum and its relatives are also smaller and less prominent, and leaves are relatively ovate, never as oblong as in the new species. Of the other pubescent- leaved Anemopaegma species, A. villosum A. Gentry, A. rugosum (Schlecht.) Sprague, A. goyazense K. Schum., and A. velutinum Mart. ex DC. have promi- nently lepidote corolla tubes. The former two also differ in possessing foliaceous pseudostipules, the latter two have more densely pubescent 3-foliolate leaves. Only two conspicuously pubescent-leaved species have glabrous corolla tubes— A. hilarianum Bureau & K. Schum. has more pubescent, smaller, ovate leaves and a paniculate inflorescence; A. brevipes S. Moore, which may be the closest relative of A. granvillei, has foliaceous pseudostipules, a shorter calyx and smaller leaflets which are much more densely puberulous beneath. Arrabidaea ornithophila A. Gentry, sp. nov.—Fic. 1. Frutex scandens; interdum consociebus glandularum in nodis inter petioles; folia tri- foliolata, foliolis elliptico-oblongis; inflorescentia floribus in panicula terminali dispositis; calyx tubulosus, bilabiatus, puberulus; corolla rosea, tubulosa; stamina subexserta, thecis pen- dulis; ovarium oblongum, lepidotum; discus annulatim pulvinatus; capsula linearis. Liana; branchlets terete, puberulous, with or without inconspicuous inter- petiolar glandular fields; pseudostipules lacking. Leaves 3-foliolate (sometimes simple in part); leaflets elliptic-oblong, rounded to acuminate at the apex, rounded at the base, 7-21 cm long, 4-10 cm wide, chartaceous, above minutely lepidote, otherwise glabrous or with a few inconspicuous trichomes along the main vein, below densely puberulous, drying dark above, light gray below; petiolules 1-4 cm long; petioles 2.5-6 cm long, lepidote and puberulous. In- florescence a terminal panicle, its branchlets densely puberulous with short glandular trichomes and longer nonglandular trichomes. Flowers with the calyx tubular, bilabiate, 10-13 mm long, 5-7 mm wide, the lobes almost 2 mm long, densely pubescent with conspicuously reddish glandular and eglandular tri- chomes; corolla cherry red, tubular, 2.3-2.5 cm long, 0.4-0.5 cm wide at the mouth of the tube, the tube 1.9-2.1 cm long, the lobes 2-3 mm long, densely puberulous outside, inside very densely pilose with exceedingly long (several mm) tangled trichomes in the upper part of the tube, these completely filling the throat, less densely pilose with shorter trichomes below, villous at the level of stamen insertion; stamens didynamous, inserted 3-4 mm from the base of the corolla tube, subexserted, the anther thecae subparallel, 3 mm long (including connective), joined for the upper 1.4-2 mm, slightly divergent basally, the con- nective extended 1.4 mm beyond the point of attachment, the filaments 1.6-1.8 cm long, the staminode 4 mm long; pistil ca. 2 cm long, the ovary oblong, taper- ing slightly towards the top, 2 mm long, 0.8 mm wide, densely glandular lepidote; disc pulvinate, 1 mm long, 2-2.5 mm wide. Fruit linear, compressed, the apex and base subacute, ca. 20 cm long, drying dark. Type: BRAZIL. PARA: Distrito Acará, Thomé Asst, Pao Vermelho, 50 m, border woods in sun, vine climbing small trees, cherry red flower, occasional, 3 Aug. 1931, Y. Mexia 6041 (MO, holotype; NY, US, isotypes). 1977] GENTRY—SOUTH AMERICAN BIGNONIACEAE d ornithophila A. Gentry.—A. Inflorescence; ren iud 6041 (MO).] x94.—B. 313 Leaves; 314 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Additional collections examined: BRAZIL. MARANHAO: Km 374 Belém-Brasilia, cipo sobre árvores, mpl 1 26 An 1960, Oliveira 3 ENSE AMAP ~ 5 Patricia, i io Jari r vermelhas, vistosas ornamental, May 1960, Froes 34938 (IAN). Rodovia Belém-Brasilia, km 92, flores vermelhas, 21 Aug. 1959, Kuhlmann & Jimbo 78 (IAN). Km 167-173 da Estrada Belém- Brasilia, flores vermelhas em cachos, 25 Apr. 1960, Oliveira 549 (IAN). Km 243- - da Rodovia Belém- Brasilia, M July 1960, Oliveira 878 (IAN). Region do Rio Jarí, estrada de Monte Pai ao Mungaba, 27 June 1968, Oliveira 4681 (IAN, NY); 2 July 1968, Oliveira 4739 (IAN, NY); 10 June 1968, Silva 2149 (IAN). Rio Jarí, Planalto de Monte Dourado, "à June 1968, sion 4548 (IAN, NY); 2 Oct. 1968, Silva 1084 (NY). Rio e serra de Monte Dourado, 140 m, 10 Nov. 1967, Oliveira 3604 (IAN, NY); 13 Nov Oliveira $534 (IAN); 18 June 1970, Silva 3227 (IAN). The closest relatives of this very distinct, presumably hummingbird-pol- linated, species are Arrabidaea trailii Sprague and Fridericia speciosa Mart. Arrabidaea trailii has a similar (but smaller) tubular red corolla with sub- exserted (but much smaller and with a minute connective) anthers and gray- drying (but never simple) leaves with densely puberulous undersurfaces. The calyx of A. trailii is smaller, subtruncate, and evenly 5-denticulate. The flower and calyx of A. ornithophila are even more similar to those of Fridericia speciosa except that the calyx of Fridericia is somewhat inflated and conspicuously 5-ridged. The anthers of Fridericia are included and the thecae divergent. The new species is so clearly intermediate between Arrabidaea trailii and F. speciosa that its existence seriously weakens the case for retention of monotypic Fridericia as a distinct genus. Cuspidaria octoptera A. Gentry, sp. nov. Frutex scandens; ramuli teretes, sine pseudostipulis, consociebus glandularum in nodis inter petiolos; folia 3-foliolata vel 2-foliolata, interdum cum cirrho, foliolis ovatis vel ellipticis, calyce cupulato, 5-denticulato, co ind 8 5 extus puberula, intus fere glabra, staminibus didynamis, eas reflexis, ovario oblongo, dense lepidoto; capsula anguste oblonga, subtetragona, 17 5 x longitudinalibus. Vine; branchlets terete, glabrous or subpuberulous, when older with scat- tered, pale, round lenticels, the interpetiolar glandular fields divided, the two halves separated by a nonglandular medial strip; pseudostipules lacking. Leaves 3-foliolate or 2-foliolate with a (presumably simple) tendril or tendril scar, never simple even at the base of branchlets; leaflets ovate to elliptic, acuminate, the base rounded, 3-8 cm long, 1.2-4 cm wide, chartaceous, the main veins plane or prominulous above, slightly raised below, somewhat puberulous along the midvein above, sparsely puberulous or pilose along the main veins below and sometimes scattered subpuberulous on the lower surface, the margins noticeably ciliate, drying brownish olive; petiolules 0.3-1.6 cm long; petioles 1.3-3.5 cm long, varyingly puberulous. Inflorescence a terminal panicle, lepidote and puberulous, bracts and bracteoles linear, to 2 mm long. Flowers with the calyx cupular, puberulous, ca. 2 mm long (with teeth), 2-3 mm wide, 5-denticulate, the teeth 0.5 mm long, extended as ribs on the outside of the calyx; corolla magenta, tubular-infundibuliform, 2.6-3 cm long, 0.9-1.3 cm wide at the mouth 1977] GENTRY—SOUTH AMERICAN BIGNONIACEAE 315 of the tube, the tube 1.8-2 cm long, the lobes ca. 0.5 cm long, puberulous out- side and on the lobes inside, the tube mostly glabrous inside, slightly glandular pubescent at the level of stamen insertion; stamens didynamous, inserted 5-6 mm above the base of the corolla, the anther thecae divaricate, pilose, reflexed forward from near the base, ca. 1.5 mm long, the blunt pilose connective ex- tended 0.3-0.4 mm; pistil 1.6-1.7 cm long, the ovary oblong, densely lepidote, 1.5 mm long, 1 mm wide; disc small, pulvinate, 0.3 mm long, 1 mm wide. Cap- sule linear-oblong, basically subtetragonal, 4-30 cm long, 1.3-2.3 cm wide in- cluding the 8 thin longitudinal wings, each wing 3-8 mm wide, glabrous, drying dark brown; seeds thin, bialate, ca. 1 cm long, ca. 3 em wide, the wings brownish- hyaline, not sharply demarcated from the seed body. Tyre: Brazit. Without locality, Nadeaud s.n. (P, holotype; MO, P, iso- types). Additional collections examined: BRAZIL. RIO DE JANEIRO: Rio Parahyba, 29 Nov. 1880, Netto et al. s.n. (R-23675, MO). Baixada Fluminense, Pilar, 30 Dec. 1939, Lutz 1565 ( R-127371, MO). Without locality, Glaziou 3769 (F). São PauLo: Without locality, Weir 516 (BM, mixed with flowering material of Arrabidaea florida DC.). This overlooked species is closely related to C. convoluta (Vell.) A. Gentry [C. pterocarpa ( Cham.) DC.] and Sandwith (in herb.) has identified flowering material (Lutz 1565) with that species, which differs most conspicuously in a merely 4-winged fruit and a larger much more deeply divided calyx with teeth 2.5-4 mm long. Vegetatively the two are extremely similar but C. octoptera differs in having uniformly 2-parted interpetiolar glandular fields (these may be 2-parted, undivided, or absent in C. convoluta), typically darker-drying leaves with noticeably ciliate leaflet margins (the leaflet margins of C. convoluta are pubescent only in var. pubescens Mello which has the whole leaf undersur- face pilose), and in the complete absence of simple leaves at the base of vege- tative shoots. Cuspidaria convoluta ranges from northern Argentina and ad- jacent Paraguay north to the states of Minas Gerais and Rio de Janeiro in Brazil where it overlaps with C. octoptera. However, the two species are probably ecologically separated since all altitudinal records for C. convoluta are from above 500 m while C. octoptera is apparently restricted to the coastal lowlands. Lundia virginalis DC. var. nitidula (DC.) A. Gentry, comb. nov. L. nitidula DC., Prodr. 9: 181. 1845. synrypes: Brazil, Minas Gerais, Martius s.n. (M, frag- ment G-DC). Brazil, Sebastianapolitana, Martius s.n. (M Bignonia nitidula Mart. ex DC., Prodr. 9: 181. 1845, nom. nud., pro syn. De Candolle (1845) was the first to systematically treat Lundia. His spe- cific concepts have proven overly narrow and four of his nine species are now generally regarded as conspecific. Bureau (1868) united L. hebantha DC. with L. virginalis DC. but maintained L. nitidula as specifically distinct. Baillon (1888) was the first to unite these two concurrently published species of de Candolle which differ only in calyx length. Since Baillon adopted L. virginalis, that name takes precedence for the species. Later Bureau & Schumann (1896- 1897) likewise concluded that L. virginalis and L. nitidula were conspecific but 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 chose to unite them under L. nitidula, treating the short-calyxed form as L. nitudula var. virginalis (DC.) Bureau & K. Schum. Since the name for the com- bined taxon must be L. virginalis under Article 57 of the International Code of Botanical Nomenclature, the new combination proposed above is needed if the long-calyxed plant is to be recognized at varietal rank. An additional problem in this complex is posed by the existence of a second short-calyxed form which dif- fers in more greenish-drying calyx and leaves, shorter, broader corolla, and white (not magenta) flower color. This has been separated as L. glazioviana Krünzl. but was considered a variant of L. virginalis by Bureau and Schumann and Sandwith (in herb.). Memora imperatoris-maximilianii (Wawra) A. Gentry, comb. nov. Bignonia 5 Wawra, Bot. Ergeb. Reise Maximilian Bras. 73, tab. 10. 1866 E: Brazil, Bahia, Wawra & Maly 156 ( W). BOE imperatoris-maximiliani (Wawra) Bureau & K. Schum. in Mart., Fl. Bras. 8(2): 279. 1897. In reviewing this species, then known only from the fragmentary type and Wawra's illustration, Sandwith (1959) noted its probable affinity with Memora rather than Pleonotoma but refrained from proposing the necessary combina- tion. Salient characters of the plant include the 5-denticulate calyx, open paniculate inflorescence, red corolla with both tube and lobes glabrous out- side, plate-shaped glands at base of the lobes, triternate leaves with rather large leaflets, and especially the terete branchlets. The latter two characters alone almost mandate placement in Memora; all other features are also consistent with this placement. Memora imperatoris-maximilianii is unusual in Memora because of its red flowers but M. magnifica ( Mart. ex DC.) Bureau has bright orange or red orange flowers and several Memora species have yellow orange flowers. The combina- tion of conspicuously 5-denticulate but otherwise truncate calyces and very minutely bracteate inflorescences are matched in Memora only by M. campicola Pilger which has a very different inflorescence, yellow flowers, and multiple compound leaves with small pubescent leaflets and M. biternata A. Samp. which has sessile leaflets and corolla lobes pubescent outside. Both of these species have the thick-foliaceous pseudostipules characteristic of most Memora species. A recent collection apparently attributable to M. imperatoris-maximilianii is now available. This is A. Lima 57-2799 (IAN) from Nazaré da Mata, Pernam- buco, and permits amplification of Wawra's description. The most noteworthy additional characteristic is the presence of conspicuous subulate pseudostipules 3-5 mm long (cf. M. cristicalyx below). The leaflets of this collection are entire to serrulate and it has the distinctly paniculate inflorescence figured by Wawra. Field notes on the Lima collection describe the flowers as "róseo nos lobos (vari- ando de róseo a lilaz bem claro) e amarelo no tubo; cálice esverd." Memora cristicalyx A. Gentry, sp. nov. Memora acutiloba Bureau, Bull. Soc. Bot. France, Mem. 58 (3f): 523. 1911, nom. nud. Habitus ignotus; ramuli teretes, glabri, sine consociebus glandularum in nodis inter peti- oles; Pr biternata, foliolis anguste ovatis, plerumque serratis; inflorescentia anguste paniculata, 1977] GENTRY—SOUTH AMERICAN BIGNONIACEAE 317 axillaris, plus minusve glabrata; flores calyce campanulato, 5-denticulato dentibus extensis in cristis, glabro, corolla tubulo-infundibuliformi, tubo extus glabro, lobis puberulis, ovario cy- lindrico dense lepidoto; fructus ignotus. Habit unknown; branchlets terete, finely striate, glabrous, drying dark with numerous round whitish lenticels, the nodes without interpetiolar glandular fields, with a raised ridge connecting opposite petioles; pseudostipules prominent, linear-subulate, 4-10 mm long. Leaves biternate, the tendril tip not seen; leaf- lets narrowly ovate, acute to acuminate, basally rounded, usually conspicuously serrate, chartaceous, 2.5-9 cm long, 1.3-4.2 cm wide, mostly glabrate, incon- spicuously scattered-lepidote, minutely subpuberulous at the base of the midvein above and below, drying blackish above, dark olive with reddish black midvein below; petiole and petiolules adaxially grooved, subpuberulous. Inflorescence a racemose axillary panicle, the lateral branches subsessile to 1.5 cm long, glabrate to subpuberulous, terminated by a pair of several mm long thinly subulate bracts, these subtending a cluster of 1 to 8 flowers on ebracteolate pedicels up to 2.5 cm long. Flowers with the calyx campanulate, subtruncate, 5-denticulate, the 0.5 mm long teeth extended as raised lateral ridges to the base of the calyx, 6-7 mm long, 5-6 mm wide, glabrous; corolla probably yellow (drying blackish yellow in type), tubular-infundibuliform, ca. 3 cm long, ca. 1 cm wide at the mouth of the tube, the tube ca. 2.5 cm long, the lobes ca. 1 cm long, the tube glabrous and the lobes puberulous outside, the lobes puberulous inside, the tube glabrous inside except at the level of stamen insertion; stamens didynamous, the filaments ca. 1.5 cm long, the anther thecae 3 mm long, somewhat divergent; ovary cylindric, 2 mm long, 1 mm wide, densely minutely lepidote; disc annular- pulvinate, 0.6 mm long, 1.5 mm wide. Fruit unknown. Type: BRAZIL. ceará: Without data, Fr. Allemáo & M. de Cysneiros 1045 ( R-127332, holotype; MO, isotype). Additional collection examined: BRAZIL. Without data, Glaziou 11232 (P, 2 sheets). This species is related to M. imperatoris-maximilianii (see above) but is other- wise remarkably isolated. Biternate leaves and terete stem mandate placement in Memora where it is the only species with serrate leaflets. Ridged calyces, rather reminiscent of Clytostoma pterocalyx Sprague, are unique in Memora and similar long subulate pseudostipules are found only in M. imperatoris- maximilianii. Memora imperatoris-maximilianii differs in an unridged glandular calyx, the corolla lobes glabrous outside and with plate-shaped glands at their bases, entire to subentire leaflets and a more open inflorescence. It is possible that additional collections from poorly known and apparently Bignoniaceae-rich northeastern Brazil will show that this and M. imperatoris-maximilianii repre- sent opposite extremes of a variable population but the available evidence sug- gests specific separation. The Paris specimens have been annotated by Bureau as “Memora acutiloba Bur., n. sp.” and that nomen nudum was used in Glaziou's "Liste des plantes du Brésil central recueillies en 1861-1895." I prefer the more descriptive epithet "cristicalyx." 318 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Tabebuia catarinensis A. Gentry, sp. nov. Frutex E 3 m altus; ur palmatim (6—)7-foliolata, foliolis Sis ellipticis vel obovatis, serra &; plabrescentibus; nflorescentia paniculata, aliquantum congesta, ramis dense s iut to- fufescentibus flores 0 tubulo-campanulato, piloso, corolla lutea, extus glabra s fauce puberula, ovario ovoideo, dense lepidoto; capsula anguste oblonga, tomentosa . barbatis, irreguliter rugulosa. Shrub 0.5-3 m tall; twigs terete, striate, minutely and glabrescently stellate- tomentose. Leaves palmately (6) 7-foliolate; leaflets oblong-elliptic to obovate, acute or very briefly acuminate, the base rounded, conspicuously and evenly serrate, the terminal leaflet to 11 cm long and 5 cm wide, the lateral leaflets pro- gressively smaller, chartaceous, when young sparsely stellate-pubescent along the main veins above and below, almost completely glabrescent at maturity, drying blackish or dark olive above and below; petiolules to 4 cm long; petioles 5-13 cm long, glabrescently stellate-tomentose. Inflorescence a several- to many- flowered, short terminal panicle, its branches densely rufescent with stellate, barbate and simple trichomes to 1 mm long, the bracts minute, subulate, to 3 mm long. Flowers with the calyx tubular-campanulate, irregularly 3-5-lobed, 12-20 mm long, 8-12 mm wide, pilose with reddish mostly barbate trichomes to 1 mm long; corolla yellow, tubular-infundibuliform, 5-7 cm long, 1.4-2.2 cm wide at the mouth of the tube, the tube 3.5-5 cm long, the lobes 1-2 cm long, drying dark brown with blackish venation, glabrous, the tube glabrous outside, inside pubescent with rather short (0.5-0.8 mm long) stiff erect trichomes de- scending from sinuses of the corolla lobes to above the level of stamen insertion, more or less glabrous at and below stamen insertion; stamens didynamous, in- serted ca. 10 mm above the base of the corolla tube, the filaments 1.7-2.2 cm long, the staminode ca. 6 mm long, the anther thecae widely divergent, 3 mm long; pistil 3.2-3.4 cm long, the ovary ovoid, ca. 2 mm long, 1.3-1.5 mm wide, densely lepidote, drying blackish, the ovules ca. 4-seriate in each locule; disc shortly cylindric, 1 mm long, 2 mm wide. Capsule linear-oblong, 5-9 cm long, 1.5-1.8 cm wide, densely reddish-brown tomentose with mostly barbate tri- chomes ca. 0.5 mm long, the surface finely and irregularly rugulose, not regu- larly striate; seeds (very immature) bialate, the wings hyaline membranaceous. Type: BRAZIL. SANTA CATARINA: Monte Crista, Garuva, campo, 750 m, ar- busto 2 m, flores amarela, 21 Oct. 1966, Klein & Ravenna 6834 (K) Additional VA Ee examined: BRAZIL. SANTA e Monte Crista, Garuva, 750 m, campo, arbusto 2 m, fruto imaturo marron, 21 Oct Klein 4» Ravenna 6828 (K); matinha, flor amarela, je Nds 3 m, 21 Oct. 1966, Klei n d ‘Ravenna 6843 (K); 900 m, campo, arbusto 0-5 m, flores amarela, 2 Sep. 1960, 5 e Klein 0 (K). Morro do Campo Alegre, Sao Francisco do Sul, 1,200 m, vu =a arbus olm, flores 11 80 3 Sep. 1960, Reitz & Klein 9766 (K); sterile, 24 Mar. 1961, zd Klein an. (K). PARANA: Mun. Campina Grande do Sul, Pico Caratuva, 1,950 m, inn do topo EA morro, flor amarela, 5 Oct. 1967, Hatsch- bach 17325 (MO). This species is superficially most similar to T. bureauvii Sandw. endemic to the vicinity of Rio de Janeiro. That species differs in being a tree to 12 m tall and in having a very shortly stellate-rufescent calyx with black-drying plate- shaped glands, a fewer-flowered, more finely tomentose inflorescence, a sparsely papillose-puberulous corolla throat, a longer (ca. 4 mm long) ovary, narrower 1977] GENTRY—SOUTH AMERICAN BIGNONIACEAE 319 leaflets, and longer, smooth-surfaced glabrous fruit. The new species was sup- posed by Sandwith & Hunt (1974) to be a form of T. chrysotricha (Mart. ex DC.) Standley. A hybrid origin from that species and T. alba (Cham.) Sandw. was also suggested as the new species is intermediate in most respects between these two species, both of which occur in Santa Catarina. Tabebuia catarinensis is ecologically distinct from lowland T. chrysotricha (below 800 m) but not from T. alba. Its flowers and inflorescence are identical to those of T. alba of which it could be a glabrescent-leaved derivative. However, the shrubby habit, shorter rough-surfaced (not striate) fruit, and uniformity of the strikingly different glabrate, rather than densely canescent, leaves support specific recognition. Hatschbach (personal communication) reports that the new species can be rather common locally at high altitudes in the Serra do Mar. LITERATURE CITED BAlLLON, H. a (1891). Histoire des Plantes. Vol. 10. L. Hachette et Cie, Paris. BunEAU, E. 1868. Révision des genres Tynanthus et Londis. Adansonia 8: 270- . SCHUMANN. 5 Bignoniaceae. In K. F. P. von Martius, Flora Bra- siliensis. Vol. 82): 1-434. Lei CANDOLLE, A. P. pk. 1845. Bodens Systematis Naturalis Regni Vegetabilis. Vol. 9. Treutel et Würtz, Paris. SANDWITH, N. Y. 1959. apu gie to the IU of Tropical America LXV. Studies in Bienoniaceae XXIV. Kew Bull. 1958: 427-4 & D. HuNr. 1974. Bignoniaceae. 7a P, R. Reitz, Flora Ilustrada Catarinense. Fasc. BIGN. Itajaí, Sta. Catarina, Brazil. NEW RECORDS OF APOCYNACEAE FOR PANAMA AND THE CHOCO' ALWYN H. GENTRY? ABSTRACT Tabernaemontana pendula and T. longipes, synonymized under T. chrysocarpa in the Flora of Panama, are recognized as distinct from it. Stemmadenia allenii was origi- nally described from fruiting and flowering material of different species, one of which— i d a, the first North American record of the South American Odontadenia cognata, and the reconfirmation of the occurrence of Fosteronia myriantha in Panama are reported. Panamanian plants referred to Tabernaemontana chrysocarpa in the Flora of Panama treatment (Nowicke, 1970) prove to represent three distinct species. These three species, somewhat similar on the basis of floral characteristics, are easily separated by vegetative and fruiting characters. Tabernaemontana pendula Woodson This species was described from a single specimen from El Valle (Allen 1734). It was compared by Woodson (1940) with T. amygdalifolia Jacq. be- cause of its exserted anthers but lumped with T. chrysocarpa, a species charac- terized by included or subexserted anthers, by Nowicke (1970) in the Flora of Panama. Tabernaemontana pendula has a much longer peduncle than either T. amygdalifolia or T. chrysocarpa. It also has wider, more elliptical leaves and a characteristically wrinkled-reticulate fruit surface. The long peduncle is also obvious in fruit. The fruit, previously undescribed, is similar in shape to that of T. chrysocarpa. Two additional collections of this species, both in fruit, are now at hand. These are Mori et al. 1912 from La Mesa (above El Valle), Coclé Province, and Mori & Kallunki 2028 from the Río Guanche area of Colón Province. Tabernaemontana longipes Donnell Smith This species was described from Costa Rica and has been thought endemic to that country. It is closely related to T. chrysocarpa and the Panamanian spe- cimens of T. longipes were included with that species in the Flora of Panama. Vegetatively T. longipes differs from T. chrysocarpa in its elliptic leaves, always broadest near the middle; the latter has narrowly obovate to oblanceolate-el- liptic leaves, broadest above the middle. The fruit of T. longipes, previously undescribed, is very distinctive with a verrucose muricate-ridged surface quite unlike the smooth, papillose or finely reticulate-ridged fruits of other Panamanian species of Tabernaemontana and Stemmadenia. I have seen no fruits of this 1 Supported by NSF grant OIP75-18202. Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. ANN. Missouni Bor. Garp. 64: 320—323. 1977. 1977] GENTRY—APOCYNACEAE 32] species from Costa Rica, but the Costa Rican and Panamanian collections seem indistinguishable on the basis of vegetative and floral characters. Tabernaemontana longipes has been collected in Panama only above El Valle de Antón, Coclé Province, where it is locally very common. It is represented by thirteen collections from El Valle in the Missouri Botanical Garden Herbarium including all the Coclé Province collections cited as T. chrysocarpa in the Flora of Panama except Allen 1734 which is T. pendula (see above). The additional collections of this species—Kennedy et al. 3035, Liesner 747, Croat 14383, Gentry & Dwyer 3612, and Gentry 6873—were all identified and distributed as T. chryso- carpa. The four Panamanian Tabernaemontana species with anthers tinged blue green can be separated by the following key: a. Anthers exserted or half-exserted; follicles narrow (more than twice as long as wide) or reniform and finely reticulate-wrinkled. Peduncle very long, exceeding the leaves; fruit reniform and finely reticulate T. pendula bb. Peduncle not 3 inflorescence not exceeding the leaves; MA narrowly elliptic, smo : quus ip aa Ies ire y or barely exserted; follicles reniform, smooth or verrucose a muricate-ridgec eaves narrowly obovate to oblanceolate 88 8 broadest above the middle; fruit smooth ; chrusocarpa cc. Leaves elliptic, broadest at the middle; fruit verrucose and muricate- ridged al T. ^ I» longipes Tabernaemontana arborea hose This species has previously been recorded from Belize to Panama. It is easily recognized by its yellow anthers inserted near the base of the corolla tube. Like many other "Central American" species, it also occurs in the northern Chocó. Two Chocó collections have been seen—Duke 12233 (MO, NY, OSU) and Duke 11169 (OSU), both from the Rio Truando. Stemmadenia allenii Woodson This species, described from E] Valle, Panama, was separated by Woodson from closely related Costa Rican S. alfari (Donnell Smith) Woodson because of its longer calyx lobes and larger corolla with broader throat and longer lobes. The type collection of S. allenii is in fruit and the fruit is of the same narrow, long-acuminate form as that of S. alfari. Many fruiting collections of S. allenii are now at hand, all with fruits of the same characteristic slender form. How- ever, a vegetatively similar species with a very different obovoid to almost orbic- ular fruit also occurs in the same wet-forest areas and the flowering material attributed to S. allenii by Woodson (1941) and Nowicke (1970) actually be- longs to the thick-fruited species (see below). The first flowering collection of the real S. allenii is Liesner 765 (MO) from El Valle, the type locality, which has the corolla throat narrower (rather than broader!) than S. alfari and calyx lobes only 4-5 mm long; in fact S. allenii proves separable from S. alfari not by the characteristics cited by Woodson but by their opposites! The real S. allenii 399 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 keys out to S. alfari in the Flora of Panama, but that species does not occur in Panama unless S. allenii itself should be considered as merely a geographic vari- ant. Stemmadenia minima A. Gentry, sp. nov. Frutex lactifer. Folia parva, anguste elliptica, acuminata, glabra. Flor calycis lobis mem- branaceis, 4-10 mm longis, corolla infundibuliformi, albida, tubo torto, staminibus tubo corollae prope mediam insertis. Folliculi crassi, obtusi, fere suborbiculati. Shrub or small tree 1.5-5 m tall; branchlets somewhat angulate, minutely papillose, lactiferous. Leaves small, narrowly elliptic, acuminate, cuneate at the base, to 11 cm long and 3.9 em wide (largest leaf 1.9-3.9 cm wide, x = 2.97 cm), glabrous above and below, membranous; petioles 2-10 mm long, not clearly differentiated from the leaf base. Inflorescence a single flower, terminal from between two dichotomous lateral branches, glabrous, with a minute triangular bracteole; calyx lobes inequal, membranous, narrowly oblong, 4-10 mm long, glabrous; corolla infundibuliform, white to cream, the tube proper 12-17 mm long, twisted 180^ at the top, the throat 15-20 mm long, the lobes obovate, ca. 1 cm long; stamens attached at the middle of the corolla tube, the anthers 4—5 mm long. Follicles thick, blunt, almost orbicular (“obovoid subreniform"), 2-3 cm long, 1.5-2.5 cm broad. Type: PANAMA. PANAMA: Cerro Jefe, 800-1,000 m, 21 Dec. 1972, Gentry 6763 (MO, holotype; duplicates were distributed as S. cf. alfari). Additional collections examined: PANAMA. CANAL ZONE: Madden Lake, Dwyer & Lallathin 7 7 5 (MO). COCLÉ: El Valle, Allen 2239, 2364; Croat 25347; Dwyer et he 4502a (all MO). coLów: Santa Rita Ridge, Gentry 6090, 6562 (both MO). PANAN Cerro Campana, Busey $61; Croat 12144; Dressler 3523; wr 8 & Kirkbride 7829A; Gentry 4934; Mori & Kallunki 1930; Porter et al. 5249 (all MO). Cerro Jefe, Croat 13028, 14444, Dressler 3333; Duke 9449 (all MO). El Llano-Cartí Road, Gentry 5071; Gentry et al. 14201, 14214; ai & Kallunki 2915 (all MO). veracuas: Mouth of Rio Concepción, Lewis et al. 2853 (MO). This species has been generally confounded with S. allenii, and, in fact, the original description of the flowers of that species are based on S. minima (see above). Discovery of the short thick fruits of S. minima prove that it is quite unrelated to S. allenii which has narrow, long-acuminate fruits. Calyx lobe length of these plants also exceeds that of S. allenii and, in fact, approaches that of S. lagunae Woodson, otherwise reported only from Bocas del Toro Province. I have previously identified collections of this entity as S. lagunae, which has a similar thick, rounded fruit. Numerous additional collections of S. minima are now at hand and it proves to differ constantly from S. lagunae in smaller leaves [largest leaf 1.9-3.9 cm wide (x = 2.97 cm) versus largest leaf 4.3-6.2 cm wide (X = 5.32 cm)] as well as shorter (x for longest lobe = 8.05 mm versus 17.2 mm), narrower, not at all imbricate calyx lobes. It usually has white flowers (some- times pale yellow or white with a yellowish center) while S. lagunae has yellow lowers (one collection reported as light yellow) and has a distinct geographic distribution. No collections are available from the critical area between Santa Fé de Veraguas where S. lagunae occurs and El Valle, the westernmost locality for S. minima, but the available evidence suggests specific recognition. 1977] GENTRY—APOCYNACEAE 393 Stemmadenia minima is fairly common in all the accessible middle elevation wet-forest areas of eastern and central Panama and is the only wet-forest species of Stemmadenia occurring east of the Canal Zone. It overlaps with S. allenii at El Valle and Cerro Campana but can be easily distinguished from that species by its very different fruit, wider corolla throat, smaller, less membranous leaves, longer calyx lobes and paler flower color. Forsteronia myriantha Donnell Smith This species was not treated in the Flora of Panama although Woodson (1935) had reported it from the Republic on the basis of a single Hayes col- lection. It has recently been recollected in Panama (Foster 4107, Barro Colo- rado Island, Canal Zone). The two Panamanian collections key to this species but have the petals sparsely pilosulose both inside and outside. Although this disagrees with Woodson's description of the petals as glabrous or very minutely papillate without, the type ( Heyde & Lux 4533 from Guatemala) also has a few long trichomes on the outside of some petals and otherwise matches the Pana- manian material. Forsteronia myriantha differs vegetatively from the other Pana- manian species in having long trichomes scattered along the leaf midvein and sometimes over the surface beneath as well as in the nerve axils. Odontadenia cognata (Stadelm.) Woodson This widespread South American species was cited from extreme eastern Pana- ma by Woodson (1935) based on a single immature collection from Puerto Obaldia, but was subsequently rejected from the Flora of Panama. Ten Panamanian col- lections are now at hand, all collected in wet-forest areas during the last few years. It is the most common species of Odontadenia on the El Llano-Cartí road and has also been collected on Santa Rita Ridge, Colón Province, and above Santa Fé, Veraguas Province. The species also reaches Costa Rica, based on Opler 1724 (MO) from La Selva, Heredia Province. Odontadenia cognata is easily told from O. puncticulosa, which it somewhat resembles, by the corolla tube ta- pering more evenly to a narrower base with the anthers inserted near the base of the tube proper rather than at the base of the throat. Several collections are noted as having pink or orange red corollas, but I have seen corollas of the more frequently reported yellow or pale yellow color only. Similar color variations occur in South America but do not appear taxonomically significant. LITERATURE CITED NowickE, J. W. 1970. Apocynaceae. In R. 4 Woodson, Jr. & R. W. Schery, Flora of Panama. Ann. Missouri Bot. Gard. 57: 59-13¢ Woopson, R. E., Foes Il Studies in the 8 IV. The American genera of Echitoideae. ouri Bot. Gard. 22: 153-306. 1940 a rei toward a Flora of Panama IV. Ann. Missouri Bot. Gard. 27: 265-364. 1941. Apocynaceae. In Contributions toward a Flora of Panama V. Ann. Mis- souri Bot. Gard. 28: 461—462 NEW TAXA AND COMBINATIONS IN ERAGROSTIS (POACEAE)! JouN T. WITHERSPOON’ ABSTRACT Eragrostis guatemalensis Witherspoon and E. intermedia Hitchc. var. appressa Wither- spoon are described as new. The former is distinguished from other North American Eragros- tis by having long pilose hairs on the lemmas y pa eas. The latter is distinguished from other members of the E. intermedia group by its short lemmas and appressed primary branchlets and pedicels. In addition, four new auae are rt . intermedia var. . H. Harvey) Witherspoon, E. intermedia var. praetermissa (L. H. Harvey) E. hirta Fourn. var. longiramea (Swallen) Witherspoon, E. trichocolea Hack. & Arech. var. floridana ( Hitche.) Witherspoon—and keys are provided to the varieties of E. interme bs E. hirta, and E. trichocolea. Eragrostis guatemalensis Witherspoon, sp. nov. cies E. intermediae varietatum intermediae et praetermissae gis sed differt ab SESS paucis pilosis propre margines lemmatum et palearum et a prima innovationibus extravaginalis, a secunda paniculis axillis dense pilosis, nervis 5 dio. inconspicuis, et caryopsibus valde sulcatus. Perennial, 65-115 cm tall. Culms erect to ascending from a somewhat knotty base, sometimes geniculate at the middle nodes; innovations extravaginal, in- frequently intravaginal, few to many, variously papillose-pilose. Leaves 4-8 per culm; sheaths slightly overlapping below, shorter than the internodes above, sparsely to densely papillose-pilose over the rounded back, along the margins and on the sides of the collar, particularly dense in the region between the margins and the “keel,” occasionally glabrate; ligules 0.3-0.4 mm long; blades linear-lanceolate, attenuate, flat to involute, 10-22 cm long, 2-5 mm wide, with dense supraligular hairs and scattered to dense papillose-pilose pubescence on the adaxial surface, glabrous to densely papillose-pilose abaxially. Panicles el- liptic to ovate, open, 24-30 cm long, 12-21 cm wide, very long exserted when mature; branches 24-30 per panicle, 10-13 cm long, ascending to spreading, curved or flexuous, floriferous 0.8-2.5 cm above the base, the lower ones soli- tary, paired or in verticels, the upper likewise and equidistant, the primary axils pilose laterally and adaxially, occasionally bearded all around the branch bases, the secondary axils often pilose laterally; primary branchlets 4-10 per branch, 30-46 mm long, ascending to spreading, capillary, mostly flexuous; secondary branchlets 0-2 per primary branchlet, mostly ascending, capillary, flexuous; pedicels 5-10 mm long ascending to spreading, capillary, flexuous. Spikelets 2-8 per primary branchlet, oblong to ovate, acute, slightly compressed, dark green to plumbeous or brownish, 3-8-flowered, 3.5-7 mm long, 1.2-2.1 mm ‘This paper is taken in part from a Ph.D. 5 submitted to the University of 5 5 5 5 would like to thank Dr. L. H. Harvey for his support through— out the study an se Hay for his assistance ae the Latin diagnoses. A grant-in-aid from Te Society 0 x m a Xi is gratefully ee 1 East 1 us waq Springfie ‘Id, Missouri 65803 ANN. Missouni Bor. Garp. 64: 324—329, 1977. 1977] WITHERSPOON—E RAGROSTIS 395 wide, the florets slightly imbricate; glumes with hyaline margins and mem- branous bodies or the first hyaline throughout, scabrous on the keels, the first ovate-lanceolate, acuminate, 1-1.5 mm long, 0.5-0.7 mm wide, the second ovate, acute, 1.5-2 mm long, 1-1.2 mm wide; lemmas ovate, acute, rounded on the back below, scabrous on the keel above, the margins and tips hyaline, the bodies membranous, often tinged with reddish purple, 2-2.4 mm long, 1.2-1.5 mm wide, the lateral nerves inconspicuous, 2-6 pilose hairs ca. 0.5 mm long in a longitudinal row on the broadest part of the lemmas between the lateral nerves and the mar- gins, the hairs fragile and caducous; paleas with hyaline margins and mem- branous bodies, usually also hyaline between the keels on the evenly convex back, ciliolate on the keels, 1.4-1.8 mm long, often with a row of hairs similar to those on the lemmas, the hairs in a longitudinal row between the keels and the margins. Caryopses oblong, strongly grooved adaxially, 0.6-0.8 mm long, 0.4-0.5 mm wide, 0.4-0.5 mm thick. Type: GUATEMALA: Alamedo, 15 July 1937, J. R. Johnston 930 (F, holo- type). 'aratypes: GUATEMALA: Solola, San Pedro, Steyermark 47100 (F, US). Guatemala, Aguilar 345 (F [EXICO. MEXICO, mi E of Ixtlhuaca, Soderstrom 515 (US). rvEBLA: El Chamizal, Ventura A. 1729 ( ENCB). This new species is similar in many respects to Eragrostis intermedia Hitchc. var. intermedia and E. intermedia var. praetermissa (proposed below). It dif- fers from the first by the hairs on the lemmas and paleas and by the pubescence on the foliage. It differs from the second by the hairs on the lemmas and paleas and by the inconspicuous lateral nerves. The only other known species of the E. intermedia group in the Western Hemisphere with hairs on the body of the lemma is the South American E. semi- nuda Trin. However, the latter is a strikingly different taxon with extremely long, villous, involute blades and a very large, diffuse panicle. The spikelets of E. seminuda are smaller and have only three or four florets. The relationships of E. seminuda seem to lie more with E. polytricha Nees and E. trichocolea Hack. & Arech. s. lat. Eragrostis guatemalensis occurs between 1,900 and 2,650 m on rocky moun- tain slopes and in oak-pine forests of central Mexico and central Guatemala. The specific epithet was derived from the taxon's occurrence in Guatemala. When first discovered among specimens from the Field Museum (F), the only known collections were those from Guatemala. Since then, however, it has been found from the states of Mexico and Puebla in Mexico. between Nicolás Romero and Progreso 1 Sohns 540 (US). de Mazapiltepec, Eragrostis intermedia Hitchc. var. appressa Witherspoon, var. nov. Varietas oreophilae affinis, sed differt lemmatibus brevioribus et pedicellis appressis. Perennial, 31-63 cm tall. Culms erect, weakly tufted, strongly anthocyanic; Leaves 2-4 per innovations intravaginal, rarely extravaginal, few, glabrous. 326 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 culm; sheaths overlapping below, slightly exceeding to slightly shorter than the internodes above, pilose along the margins and on the collar sides and throat; ligules 0.2-0.3 mm long; blades linear-lanceolate, acuminate-attenuate, flat or involute below, tightly involute above, appressed or ascending, 2-9 cm long, 1-2 mm wide, densely pilose above the ligule and a short way up the adaxial surface. Panicles elliptic when young, mostly deltoid when mature, open to somewhat condensed, stiff, straight or curved at the tip, 12-24 cm long, 5-20 cm wide, short to long exserted; branches 10-24 per panicle, 5-13 cm long, ap- pressed-ascending to ascending-spreading, infrequently spreading, straight, florif- erous 0.9-3.1 cm above the base, the primary and secondary axils pilose laterally and abaxially, occasionally also adaxially; primary branchlets 0-9 per branch, 13-45 mm long, appressed to ascending, capillary; secondary branchlets absent; pedicels 1-6 mm long, appressed to appressed-ascending, capillary. Spikelets 1-6 per primary branchlet, ovate, acute, barely compressed, reddish purple, 3-5- flowered, 2.4-3.7 mm long, 1.2-1.6 mm wide, the florets slightly imbricate, tightly so when young; glumes hyaline or the second with hyaline margins and a mem- branous body, strongly scabrous on the keels, the first lanceolate to ovate, acu- minate to acute, 1-1.5 mm long, 0.5-0.6 mm wide, the second ovate, acute, 1.3-1.6 mm long, 0.7-0.8 mm wide; lemmas ovate to broadly so, acute, rounded or slightly keeled below, scabrous on the keel above, the tips hyaline, the bodies membranous, often tinged with reddish purple, 1.5-1.7 mm long, 1-1.4 mm wide, the lateral nerves very weak; paleas hyaline, ciliolate on the keels, 1.2-1.5 mm long. Caryopses oblong to quadrate, weakly grooved adaxially, reticulate, 0.5-0.7 mm long, 0.3-0.5 mm wide, 0.3-0.5 mm thick. Type: Mexico, Jalisco: 7 mi S of Zacatecas-Jalisco border, 11 Oct. 1972, Harvey & Witherspoon 9344 (US, holotype; ENCB, MO, MONTU, NY, RM, TAES, W, isotypes). This taxon is phenetically close to Eragrostis intermedia var. oreophila (pro- posed below) but is distinguished from it by its shorter lemmas and appressed pedicels. This variety is known only from the type locality. The area is a roadside swale containing limestone, at approximately 1,830 m. The vegetation is sparse and the plants of var. appressa grow in small tufts scattered throughout the area. Oaks and junipers are common on the margins of the swale. The plants are a striking deep-red color and upon emergence from the sheath, the panicles appear spikelike, resembling a Sporobolus or Muhlenbergia with narrow, compact panicles. All orders of branching remain appressed for some time following emergence, but even after the main branches spread, the pri- mary branchlets and pedicels remain appressed, hence the derivation of the name. Eragrostis intermedia Hitchc. var. oreophila (L. H. Harvey) Witherspoon, comb. et stat. nov. Eragrostis oreophila L. H. Harvey, Bull. Torrey Bot. Club 81: 407. 1954. rype: Mexi idalgo, Jacala, stony mountain side, 4,500 ft, 29 June 1939, V. H. Chase 7223% (US. holotype; ARIZ, GH, MICH, MO, TEX, isotypes). 1977] WITHERSPOON—ERAGROSTIS 2 i -1 This variety occurs primarily on mountain slopes in central Mexico. It grows in fairly deep, sandy-clay soils at elevations above 1,380 m. It is not a common taxon in most of its range, but seems to be quite abundant in the mountains around Jacala, Hidalgo. It is known from single collections in both Baja California and Nuevo León. The wide separation of these collections may reflect inadequate sampling but it may be an indication that the taxon has arisen more than once. Variety intermedia is common in all these areas. How- ever, the two taxa remain relatively distinct in these areas of sympatry. One collection from Chihuahua (Gentry 8157, Sierra Charuco, MICH, UC, US) has pubescent foliage not characteristic of other known specimens. Eragrostis intermedia Hitchc. var. praetermissa (L. H. Harvey) Witherspoon, comb. et stat. nov. Eragrostis praetermissa L. H. Harvey, Bull. Torrey Bot. Club 81: 408. 1954. Type: Guate- mala, Dept. Baja Verapaz, Santa Rosa, July 1887, H. von Türckheim 1292 (US, holo- type). Eragrostis intermedia var. praetermissa occurs at high elevations in Mexico and Central America. It grows primarily in the pine-oak zone in deep, loamy soils. It is not common in any part of its range but may be more prevalent than herbarium material indicates. Since it grows in seemingly native vegetation, most collectors would probably not encounter it if they botanize mostly along roads and established trails. This variety, although more common than var. oreophila, is perhaps not as distinct. Intermediates between var. intermedia and var. praetermissa have been noted rarely, but var. praetermissa can usually be distinguished by its extra- vaginal buds and papillose-pilose sheaths. The following key will effectively separate the varieties of Eragrostis inter- media. a. Buds extravaginal; at least the lower sheaths densely papillose-pilose; blades usually flat, often exceeding 5 mm wide; spikelets dark green, the lemmas with strong lateral nerves, rarely weak pare var. praetermissa aa. sor ee 1 dur sheaths ¿lubos xen iar a [tut " pee at the apex or ith scattered hairs; blades mostly involute, usually less than 5 mm wide; spikelets light green to uia often reddish, the lemmas with weak to inconspicuous ateral nerves, rarely strong — b. Panicles duesdnmound, with = branchlets at least on the lower primary branchlets, averaging more than 6 primary branchlets per branch; leaves few to many, not "crowded at the base; n generally equal to or shorter than the spike | A ² GG RR AMONG var. intermedia bb. Panicles Los secondary branchlets, usually with less idus an a primary bran s per br ənch; leaves few, crowded at the base; pedicels generally longer 5 the spikelets pencils and pedicels appressed to the main branches; lemmas less than CC1%%%%%%CCC ̃ͤ ͤ A var. dnx Branchlets and pedicels ascending to pedal. pene is pers than equal to 1.8 mm long — ay . e 2 e 328 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Eragrostis hirta Fourn. var. longiramea (Swallen) Witherspoon, comb. et stat. nov. Eragrostis longiramea Swallen, J. Wash. Acad. Sci. 21: 437. 1931. Type: Mexico, Tamaulipas San Carlos, Pico del Diablo, vic. Marmolejo, 12 Aug. 1930, H. H. Bartlett 10910 (US, holotype; GH, LHH,* MICH, isotypes). This variety occurs in dry, rocky soils along forest borders and streams from 900 to 3,500 m. Its distribution is generally more northern than typical var. hirta. It is known only from the mountains of Tamaulipas, Nuevo León, and San Luis Potosí in Mexico. Variety longiramea is much larger than var. hirta, often approaching 2 m in height. The two varieties are quite similar vegetatively, var. longiramea usually having more flattened, longer and wider blades. However, they are easily dis- tinguished by their panicles, as shown in the following key. a. Panicle less 1 5 or equal to 45 em long and 10 em wide, the branches less than or 1 lo equal to 10 cm long — . var. hirta aa. Panicle 3 if Pta 50 em long and 18 em wide, the branches greater than 15 c — eee aaa eee jj te 55 ane Eragrostis trichocolea Hack. & Arech. var. floridana (Hitchc.) Witherspoon, comb. et stat. nov. Eragrostis floridana Hitchc., Amer. J. Bot. 2: 308. 1915. Type: United States, Florida, dry woods near Ta ampa, Oct. 1885, ^a H. Curtis 3494 [US, holotype; BR, F, ISC, LE, MO, NY (2), PH, TAES, TENN, US (2), 5 s]. HORE purpusii Jedw., Bot. Arch. 5: 201. 1924. rype: Mexico, Puebla, Cerro de Gavilan, Aug. 1909, C. A. Purpus 4084 (UC, lectotype, "here designate d; US, fragment of holo- ty x B). The holotype was destroyed in Berlin; the US fragment consists only of a small branch with a few spikelet Eragrostis trichocolea var. floridana is found in sandy pinelands in coastal and central Florida and in sandy ground near Huntsville, Texas. It also occurs in sandy ground and sandy pine forests in central Mexico and Central America at higher elevations. This variety is distinguished by its smaller lemmas, more pubescent foliage and its distribution. Hitchcock described E. floridana in 1915 based on collections from Florida and Orizaba, Mexico. However, after seeing material of E. trichocolea in Euro- pean herbaria, he decided that E. floridana was conspecific with that taxon. From notes on the type specimen of E. floridana at US, he apparently made this change in 1918 and the taxon was treated as E. trichocolea in both editions of Manual of the Grasses of the United States (1935 and 1950). However, Hitch- cock had not seen the holotype of E. trichocolea that is housed in Montevideo (MVM). The “types” seen by Hitchcock and most other North American work- ers were fragments of two Arechavaleta collections procured by Agnes Chase in 1922 and placed in the U.S. National Herbarium. She obtained these frag- ments from the Hackel Herbarium in Vienna and the labels on those specimens do not match the label on the holotype. Therefore, the fragments at US do not aaa types, although they are, in fact, E. trichocolea var. trichocolea. * LHH denotes the personal herbarium of Dr. L. H, Harvey, University of Montana. 1977] WITHERSPOON—ERAGROSTIS 329 After examining the holotype and other South American material, I feel that the North and Central American specimens are phenetically distinct at the vari- etal level. Eragrostis purpusii falls within the morphological limits of E. trichocolea var. floridana. The varieties of Eragrostis trichocolea may be distinguished by the following Lemmas greater than or equal to 1.8 mm long and 1.4 mm wide, averaging over 2 mm in length; blades variously papillose-pilose less than 1⁄4 thea length; South Amer S var. ae aa. 5 less than or equal to 1.8 mm long and 1.3 mm wide, avera ging less yea 1.7 mm in length; blades densely iit a at least 74 their EM 19 ar Central America go = REALIGNMENT OF THE SPECIES PLACED IN EXOGONIUM (CONVOLVULACEAE )' DANIEL F. AUSTIN? Exogonium has never been widely accepted as a genus although there have been proponents of this rank since Choisy (1834, 1838, 1845) first described the taxon. House (1908), Matuda (1963) and Standley & Williams (1970) have been among the recent authors keeping the species as a separate genus. Others have suggested that the species could better be ranked at some infrageneric level. Grisebach (1864) reduced it to a section of Ipomoea, while Meisner (1869) considered the plants a subgenus. Since the origin of the name Exogonium by Choisy (1834) 31 species have been placed in the taxon, many authors varying the definition of the group slightly. Although usually unstated, the major criteria for inclusion in the taxon were red flowers, salverform corollas, and exserted stamens and stigmas. Floral morphology suggests that the species included in Exogonium are mostly adapted for bird pollination; the species exhibit the characters classically associated with this syndrome (van der Pijl, 1960, 1961; Meeuse, 1961; Percival, 1965; Faegri & van der Pijl, 1971). However, a polyphyletic taxon has been created because species from several lines have been lumped solely on the basis of a common pollination system. e following treatment is a revision of the binomials placed in Exogonium. Several other related species are also included. Some of the nomenclatural and biological problems within Ipomoea have been discussed elsewhere (Verd— court, 1957, 1963; Austin, 1975a, 1975b). Consideration of all morphological criteria indicates that the species pro- posed for inclusion in Exogonium should be placed in the following taxonomic groups. Group 1. Exogonium velutifolium House [Bull. Torrey Bot. Club 35: 100. 1908. Type: Mexico, Oaxaca, Nelson 1877 (GH, holotype; US, isotype)] is not a member of the Convolvulaceae but the Acanthaceae. The correct name is Ruellia velutifolia (House) Wasshausen & Austin (Phytologia 25: 433-437 1973 ‘This study was supported by grants from the Division of Sponsored Research, Florida Atlantic University, and a grant from the National Science Foundation through the Smithson- ian Institution ( Summer Institute in Systematics—V. Species 5 specimens in the herbaria at A, FAU, x G, GH, IJ, K, L, LE, M, MEXU, MO, NY, US, and W were examined during visits and on loan. Exc cept w where p are cited from other herbaria, material from these Vi deri formed the basis of the stuc My thanks are ex- tended to yaqa and staff of the institutions cited. K. R. Robertson Arnold Arboretum ) has revised the us V and provided especially useful comments. W. G. D’Arcy ( Missouri ind i 'Garde n) criticized the original manuscri Department of Biological Sciences, Florida Atlantic University, Boca Raton, Florida 33431. ANN. Missouni Bor. Garb. 64: 330-339. 1977. 1977] AUSTIN—REALIGNMENT OF EXOGONIUM 331 Group 2. Exogonium filiforme (Desr.) Choisy is the only member of the genus Jacquemontia with bird-pollinated flowers. The small seeds and capsule which breaks into several sections at dehiscence, among other characteristics, support the conclusion that this is a Jacquemontia (see Robertson, 1971). Jacquemontia solanifolia (L.) Hall. f., Bot. Jahrb. Syst. 16: 542. 1893. Basionym: Ipomoea solanifolia L., Bp. Pl. 161. 1753. Lecrorype: Ipomoea foliis cordatis Plumier, Cat. Pl. Amer., p. 3 in Nova Pl. Amer. Gen. Tab. XCIV. fig. Synonyms: Quamoclit solanifolia (L.) Choisy in Meta Prodr. 9: 335. 1845. Exogonium solantjolium (L.) Britton, Mem. Brooklyn ae en 1: 1918. omoea filiformis Jacq., Enum. Pl. Carib. n ; Sel. Stirp. Amer. 27, pl. I9. 1763. LECTOTYPE: illustration by Jacquin, i E^ TA e d filiformis (Jacq.) Desr. in Lam., Encycl. Méth. Bot. 3: 555. 1789. Exogonium filiforme (Jacq.) Choisy, Mem. Soc. Phys. Genéve 8: 129. 1838 — Distribution: Puerto Rico, Guadalupe, Antigua, St. Barthelemy, St. Croix, St. Thomas, Martinique, Tortola. Group 3. House included Exogonium racemosum, E. wrightii and E. ru- dolphii in his concept of the genus. These three names have been reduced to two species and placed in Turbina by O'Donell (1960) and Liogier (1968). The accrescent sepals indicate that this is the proper disposition. A new com- bination does need to be made. 1. Turbina racemosa ( Poir.) D. Austin, comb. nov. Basionym: Ipomoea racemosa Poir. in Lam., Encycl. Méth. Bot. Suppl. 4: 633. 1816, non Roth, 1821, nec Griseb., 1866. tyre: St. Domingo. Possibly in LAM-P, not in the microfiche. Interpretation based on the protologue. Synonyms: Convolvulus racemosus (Poir.) Sprengel, Syst. Veg. 1: 600. 1825. Exo- gonium racemosum (Poir.) Choisy, Mem. Soc. Phys. Genéve 8: 128. 1838. Calystegia berterii Sprengel ex Hall. f., Bot. Jahrb. Syst. 16: 558. 1893, nom. nud., pro syn. Ipomoea bracteata Rudol. ex Ledeb. in ME Neues J. Bot. 2: 292. 1807, non Cav., 1799. rype: St. Domingo, Rudolphi nier at E). Ipomoea de Roem. & Schult., Syst. Veg. 4: 222. 1819. Pharbitis bracteata ec B Ledeb.) Choisy in DC., Prodr. 9: 344. 1845. Rivea bracteata (Rudol. ex Ledeb.) Hall. pe D Syst. 18: 158. 1894. Turbina rudolphii (Roem. & Schult.) O'Donell, wg T 64 Convolvulus altissimus Sprengel, Syst. Veg. 613. ns TYPE: Hispaniola, Bertero ( MO, SET. Ipomoea altissima ( Sprengel) Es ex G. Don, Gen. Syst. 4: 273. 1838. Distribution: Cuba, Haiti, Dominican Republic. 2. Turbina wrightii (House) Alain, Brittonia 20: 152. 1968. Basionym: Exogonium wrightii House, Bull. Torrey Bot. Club 35: 99. 1908. TYPE: Cuba, Wright 1650 (GH, holotype; MO, isotype). Synonyms: Ipomoea wrightii (House) Alain, Mem. Soc. Cub. Hist. Nat. "Felipe Poey" 1955. 22: 123. Ipomoea racemosa sensu Griseb., Cat. Pl. Cub. 205. 1866, non Poir., 1816. Distribution: Endemic to Cuba. 332 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Group 4. Three binomials illustrated by Sessé & Moçiño have been placed in Ipomoea sect. Quamoclit by Choisy (1845) and O'Donell (1959). Their caudate sepals clearly indicate affinity with that species group. 1. Ipomoea hastigera H.B.K., Nov. Gen. Sp. Pl. 3: 87. 1819; O'Donell, Lilloa 99: 49. 1959 Exogonium curviflorum Sessé & Mocino ex Choisy in DC., Prodr. 9: 336. 1845, nom. pro syn. 2. Ipomoea neei (Sprengel) O'Donell, Lilloa 29: 69. 1959. Exogonium umbellatum Mocino & Sessé ex Choisy in DC., Prodr. 9: 336. 1845, nom. pro syn. 3. Ipomoea neei (Sprengel) O'Donell, Lilloa 29: 69. 1959. Exogonium corimbosum Sessé & Mocino ex O'Donell, Lilloa 29: 71. 1959, nom. pro syn. A fourth species was not included in O’Donell’s discussion of the Quamoclit group. The species is an Ipomoea; the flowers appear to be bird pollinated; and the morphology indicates affinity with the Quamoclit group. 4. Ipomoea uhdeana (Fenzl. ex Hall. f.) D. Austin, comb. nov. Basionym: Exogonium uhdeanum Fenzl. ex Hall. f., Bot. Jahrb. Syst. 16: 9. 1893. A new name for Quamoclit tubulosa Mart. & Gal. Synonyms: Quamoclit tubulosa Mart. & Gal, Bull. Acad. Roy. Sci. Bruxelles 12: 270. 1845. rvrk: Galeotti 1393 (W, isotype). Ipomoea un (Mart. & Gal.) Hemsl., Biol. Centr. Amer., Bot. 2: 395. 1882, non Roem. & Schult., 181€ Group 5. Ipomoea bracteata, the type species of Exogonium, shares many characteristics with I. purga and its allies. Only three of the nine species in the alliance have been placed in Exogonium. Ipomoea: Exogonium alliance. Exogonium Choisy, Mem. Soc. Phys. Genève 6: 443. 1834 Ipomoea subgen. Exogonium (Choisy) Meisn. in Mart., FL Bras. 7: 221. 1869. Ipomoea sect. Batatas subsect. Emeticae House, Ann. New York Acad. Sci. 18: 239. 1908. l. Ipomoea bracteata Cav., Icon. Descr. Pl. 5: 51, pl. 477. 1799. type: Cited as Ipomoea ?bracteata by Cav.; based on two collections: Mexico, “Mazatlan duabus leucis" Sessé & Mogifio and "quattuor a Chipalcingo” Sessé & Mo- çiño (probably MA, not seen). Interpretation based on plate 477 by Cavanilles. Syr ns: ar 8 cincta Roem. & Schult., Syst. Veg. 4: 254. 1819, nom. illeg. Exo- dies pe teatum ( Cav.) Choisy, Mem. Soc `. Phys. Genève 6: 443. 1834. omoea spicata H.B.K. Nov. Gen. Sp. Pl. 3: 112. 1819. TYPE: Mexico, oe d> Bonpland i not seen, microfiche seen). Exogonium spicatum (H.B.K.) Choisy, Mem. Soc. Phys. G e 8: 128. 1838, nom. illeg. Eun olivae Bárcena, Viaje C av. Cacahuam. 29. 1874. TYPE: Mexico, RM ip^ m bly MEXU, not found). 5 based on the Bärcena pla volvulus bractiflorus Sessé & Mog Pl. . Hisp. 38. 1887; n. Now. "Hisp. 22. 1893. rype: Mexico, Sessé Moçiño fon balls us "Their Icones 207 i ed in the 1893 publication, but I have not been able to obtain a copy of this. The 5 used here is based on the protologue. - AUSTIN—REALIGNMENT OF EXOGONIUM 333 Ke} -1 — Ipomoea bracteata var. pubescens Robinson & Greenman, Amer. J. Sci. 50: 160. 1895. TYPE: Mexico, Jalisco, Pringle 4734 (MO, holotype ). Exogonium bracteatum var. pubescens (Robinson & Greenman ) House, Bull. Torrey Bot. Club 35: 101. 1908. Distribution: Baja California, Jalisco, Sinaloa, Oaxaca, Tepic, Morelos, So- nora, Chihuahua, Michoacan, and Guerrero (Mexico). 2. Ipomoea dumosa ( Benth.) L. O. Williams, Fieldiana, Bot. 32: 190. 1970. Basionym: Exogonium dumosum Benth., Pl. Hartw. 46. 1840. type: Hart- weg s.n. (K, holotype, not seen; F, photo). Distribution: Mexico. This species is very close to I. purga and has been considered synonymous with that population by some. 3. Ipomoea elongata Choisy in DC., Prodr. 9: 355. 1845. rype: Mexico, Oaxaca, Andrieux 212 ( G-DC, holotype; US, photo). Distribution: Mexico. See Matuda, Anales Inst. Biol. Univ. Nac. México 35: 75. 1964 4. Ipomoea emetica Choisy in DC., Prodr. 9: 376. 1845. rype: Mexico. Based on an unpublished plate by Sessé & Moçiño (not found). Synonyms: Ipomoea sagittata Sessé & Mocino ex Choisy in DC., Prodr. 9: 376. 1845, nom. 55 syn., non Poir. )omoea caudata Fernald. Proc. Amer. Acad. Arts 36: 498. 1901. TYPE: Mexico, Morelos, Pringle 8448 (GH, holotype). Distribution: Mexico. See Matuda, Anales Inst. Biol. Univ. Nac. México 36: 85. 1965 5. Ipomoea hintonii L. O. Williams, Econ. Bot. 24: 400. 1970. rype: Mexico, Hinton et al. 8474 (F, holotype). Distribution: Mexico. 6. Ipomoea purga (Wender.) Hayne, Arzneigewüchse 12: tab. 33, 34. 1833. Basionym: Convolvulus purga Wender., Pharm. Central-Blatt 1: 457. 1830. TYPE: Based on plants grown from seed collected in Mexico by Schiede, prob- ably not preserved. Interpretation based on Hayne plates. Synonyms: Exogonium te dicor i en Pl. 5 artw. 46. 1840. 5 schiedeana Zucc., Flor 1831. PE: Based on plants collected in Mexico by Schiede and Pere d by B Probably based on the same Schiede col- lections as I. purga. Distribution: Mexico, Guatemala, El Salvador, Honduras, Costa Rica, Pan- ama. 7. Ipomoea seducta House, Ann. New York Acad. Sci. 18: 241. 1908. TYPE: Guatemala, Alta Verapaz, Tuerckheim 7926 (GH, US, isotypes). Distribution: Mexico, Guatemala. See Standley & Williams, Fieldiana, Bot. 24 (9):51. 1970. 8. Ipomoea suffulta ( H.B.K.) G. Don, Gen. Syst. 4: 276. 1838. Basionym: Convolvulus suffultus H.B.K., Nov. Gen. Sp. Pl. 3: 102, pl. 211. 334 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 1819. tyre: Mexico, Volcán de Jorullo, Humboldt & Bonpland (P, not seen, microfiche seen). 9. Ipomoea urbinei House, Muhlenbergia 3: 41, pl. 2. fig. b. 1907. Type: Mex- ico, Volcán de Colima, Bárcena 214 (presumably MEXU, not found). Distribution: Known only from the type collection. See Matuda, Anales Inst. Biol. Univ. Nac. México 35: 67. 1964 Group 6. The species in this group form a relatively homogeneous assem- blage within Eriospermum which is normally recognized as a section or sub- genus of Ipomoea (Verdcourt, 1963; Austin 1975a). Comose seeds separate these species from the others that have been placed in Exogonium. Some justifica- tion for the separation of this group from Eriospermum could be made. Most of the species listed here are adapted for bird pollination while others in Erio- spermum are bee pollinated. If the two groups were placed in different taxa, several species-pairs (e.g., I. eggersii/I. steudelii and I. viridiflora/I. carolina) would be separated. While closely related, one species of these pairs conforms to the bee-pollination syndrome (I. eggersii), and the other to the bird-pollina- tion syndrome (I. steudelii). If this alliance were separated from Eriospermum, the closely related species of the pairs would be placed in separate taxa. Ipomoea mirandina/I. microdactyla alliance. Exogonium Choisy, Mem. Soc. Phys. Genéve 6: 443. 1834, in part, excl. type species. Ipomoea sect. Exogonium (Choisy) Griseb., Fl. Brit. W. I. 472. 1862, excl. type species. 1. Ipomoea argentifolia A. Rich. in Sagra, Hist. Cuba 11: 131. 1850. TYPE: Isle of Pines, Richard (P, not seen). re d Exogonium argentifolium (A. Rich.) House, Bull Torrey Bot. Club 35: 102. 1908. Ipomoea praecox Wright in Sauv., Fl. Cubana 107. 1873; Anales Acad. Ci. Méd. Habana 7: 46. 1870. type: Wright 3644 (US, isotype). Distribution: Cuba, Isle of Pines, Mexico (Oaxaca, Puebla). Although I have not seen the type of Richard's species, it is the only popula- tion on the Isle of Pines matching the protologue. The distribution of this spe- cies is worthy of note in that the plants are disjunct from western Cuba to the western slopes of the Sierra Madre Occidental in Mexico. The species is prob- ably native to Cuba and not introduced since the allied species Ipomoea lachnea Sprengel ( Bertero, MO, isotype) occurs in the Dominican Republic. 2. Ipomoea carolina L., Sp. Pl. 160. 1753. Type: Based on illustration in Catesby, Nat. Hist. Carolina 2: 9, tab. 91 (lectotype). Synonyms: Exogonium pedatum Cholsy, Mem. Soc. Phys. Genève 8: 130. 1838. TYPE: Santo Dam ags. Poiteau ( G- , not seer PIpomoea piaua Rudolphi ex. Le de b. & Adlerstam, Pl. Doming. 14. 1805; Ledeb., Neues J. Bot. 2: 292. 1807. type: Rudolphi coll.? (not in LE). Distribution: Haiti, Dominican Republic, Bahama Islands. The synonymy listed here is based entirely on the descriptions since the types have not been seen. The protologue of I. clausa is general enough that one can- not be sure of the species. Some have thought that this name applies to I. triloba 1977] AUSTIN—REALIGNMENT OF EXOGONIUM 335 (Matuda, 1965: 100). Perhaps I. clausa does apply to that species, but the de- scription appears to me to better fit the population treated here as I. carolina. While I was in Geneva I looked for the type of E. pedatum but did not find it. Additional searching will perhaps solve that problem of synonymy. 3. Ipomoea conzattii Greenman, Publ. Field Columbian Mus., Bot. Ser. 2: 258. 1907. Type: Mexico, Conzatti 1666 (F, holotype). Synonym: Exogonium conzattii (Greenman) House, Bull. Torrey Bot. Club 35: 102. 1908. Distribution: Mexico (Guerrero). m. Ipomoea concolora (Matuda) D. Austin, comb. nov. Basionym: Exogonium concolorum Matuda, Anales Inst. Biol. Univ. Nac. Mexico 36: 116. 1965 (1966). Type: Kruse 844 (MEXU, holotype ). 5. Ipomoea clarensis Alain, Mem. Soc. Cub. Hist. Nat. “Felipe Poey” 22: 121 1955. rype: Cuba, Leon & Roca 7959 (NY, holotype, not found). Although the type was not found, there is a specimen [Howard 6565 (US)] annotated by Alain. Distribution: Endemic to Cuba. This species is very similar to I. micro- dactyla. The main difference is the flower color: red in I. microdactyla and white in I. clarensis. 6. Ipomoea cubensis (House) Urban, Symb. Antil. 9: 427. 1925. Basionym: Exogonium cubense House, Bull. Torrey Bot. Club 35: 105. 1908. TYPE: Britton & Shaffer 495 (NY, holotype). 7. Ipomoea desrousseauxii Steud., Nom. Bot, ed. 2. 816. 1841. Type: Based on Convolvulus eriospermus Desr. Basionym: Convolvulus eriospermus Desr. in Lam., Encycl. Méth. Bot. 3: 967. 1789. tyre: probably in P-LAM, not seen. Synonyms: Exogonium eriospermum ( Desr.) room Mem. Soc. Phys. Genéve 8: 130. 1838. Nx eriosperma (Desr.) Raf., Fl. Tell. 4: 1838, non Beauv., 1807. — The specimen in the DeCandolle herbarium [Santo Domingo, Bertero s.n. (G DC)] matches the description of Desrousseaux and the synonymy is based on that specimen. Distribution: Santo Domingo. 8. Ipomoea eggersii (House) D. Austin, comb. nov. Basionym: Exogonium eggersii House, Bull. Torrey Bot. Club 35: 104. 1908. TYPE: St. Thomas, Feb. 1887, Eggers (NY, holotype, not found; G, L, isotypes). Distribution: St. Thomas, Tortola. This species has been confused with I. steudelii. Differences between them are few; the major distinction is that I. eggersii has white bee-pollinated flowers and I. steudelii has red bird-pollinated flowers. 9. Ipomoea fuchsioides Griseb., Cat. Pl. Cub. 205. 1886. type: Cuba, Wright 3095 (MO, isotype). 336 ANNALS OF THE MISSOURI BOTANICAL GARDEN Vor. 64 Synonyms: Exogonium fuchsioides (Griseb.) House, Bull. Torrey Bot. Club 35: 101. 1908 Distribution: Endemic to Cuba. 10. Ipomoea incerta (Britton) Urban, Symb. Antil. 9: 247. 1924. Basionym: Exogonium incertum Britton, Mem. Torrey Bot. Club 16: 94. 1990. rype: Cuba, Shafer 1235 (NY, holotype). Distribution: Endemic to Cuba. ll. Ipomoea jalapoides Griseb., Cat. Pl. Cub. 202. 1886. TYPE: Cuba, Wright 3097 ( MO, isotype). Synonym: Exogonium jalapoides (Griseb.) House, Bull. Torrey Bot. Club 35: 101. 1908. Distribution: Endemic to Cuba. 19. Ipomoea longistaminea O'Donell, Lilloa 23: 488. 1950. type: Brasil, Bahia, Rose & Russell 19784 (US, holotype). Distribution: Endemic to Brasil and apparently to the state of Bahia. 13. Ipomoea leuconeura Urban, Symb. Antil. 3: 350. 1902. svwTYPES: Haiti, Ehrenberg 134 (US, fragment), Picarda 16 (US, fragment), Buch 5 (not seen). Synonym: Exogonium leuconeurum (Urban) House, Bull. Torrey Bot. Club 35: 106. 908. Distribution: Endemic to Haiti. 14. Ipomoea microdactyla Griseb., Cat. Pl. Cub. 204. 1886. TYPE: Cuba, Wright 3094 (MO, isotype). Synonyms: Exogonium microdactylum (Griseb.) House, Bull. Torrey Bot. Club 35: 102. Exogonium microdactylum, var. nem House, Bull. Torrey Bot. Club 35: 103. 1908. Cuba, Wright 3102 (MO, is inopes repanda sensu 8 pod Pl Cub. 204. 1886, non Jacq. 1760. Distribution: Florida, Bahamas, Cuba. These vines are found only in the rocky pinelands of Dade and Monroe counties in Florida. In the Bahamas (Inagua) the Bahama Woodstar hummingbird visits and pollinates the flowers. 15. Ipomoea mirandina (Pittier) O'Donell, Lilloa 26: 370. 1953. Basionym: Exogonium mirandinum Pittier, J. Wash. iue Sci. 21: 143. 1931. TYPE: Venezuela, Pittier 12217 (VEN, holotype; US, isot Distribution: Known from Venezuela and Panama; “andadi in Co- lombia also but no specimens seen. 16. Ipomoea repanda Jacq., Enum. Pl. Carib. 13. 1760; Sel. Stirp. Amer. 28, pl. 20. 1763, non Griseb., 1886. Lecroryre: illustration by Jacquin, pl. 20. 1763. Synonyms: Convolvulus repandus ( Jacq.) Desr. in Lam. , Encycl. Méth. Bot. 3: 555. 1789. an Á repandum (Jacq.) Choisy, Mem. Soc. Phys. Gen RM 8: 198. 1838 Distribution: Puerto Rico, Tortola, Cuba, Barbuda, Antigua, Martinique, Dominica, Guadeloupe, St. Vincent, Montserrat, St. Lucia, St. Jan. 1977] AUSTIN—REALIGNMENT OF EXOGONIUM 337 17. Ipomoea retropilosa (Pittier) D. Austin, comb. nov. Basionym: Exogonium retropilosum Pittier, J. Wash. Acad. Sci. 21: 143. 1931. type: Venezuela, Mérida, Pittier 12698 (VEN, holotype; MO, US, isotypes). Distribution: Endemic to the coastal mountains of Venezuela. 18. Ipomoea shinnersii D. Austin, nom. nov. Basionym: Exogonium luteum House, Bull. Torrey Bot. Club 35: 103. 1908. TYPE: Mexico, Conzatti & Gonzalez 668 (GH, holotype; NY, isotype). Because of Ipomoea lutea Hemsley (Diagn. Pl. Nov. 34, tab. 60. 1878) a new name is required for House’s species. The new name commemorates the late Lloyd Shinners, a student of Convolvulaceae. Distribution: Mexico. (Guerrero), 19. Ipomoea shinnersii var. woronovii (Standley) D. Austin, comb. et stat. nov. Basionym: Exogonium woronovii Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 171. 1932. Type: Mexico, Woronow 2906 (F, holotype). Distribution: Mexico (Michoacán ). 20. Ipomoea signata House, Muhlenbergia 3: 46. 1907. TYPE: Guatemala, Nelson 3595 (US, holotype). Distribution: Mexico, Guatemala, Venezuela. See Matuda, Anales Inst. Biol. Univ. Nac. México 35: 72. 1964; Standley & Williams, Fieldiana, Bot. 94 (9): 53. 1970. The distribution of this species is unusual in that it appears to be absent from a large part of Central America and reappears in the coastal mountains of Venezuela. 2]. Ipomoea steudelii Millsp., Publ. Field Columbian Mus. Nat. Hist., Bot. Ser. 2: 86. 1901. Type: Based on Ipomoea arenaria (Choisy) Steud. Synonyms: Exogonium arenarium Choisy, Mem. Soc. Phys. Genéve 8: 129, pl. 1. : TYPE: Puerto Rico, Bertero (G-DC, lectotype). Ipomoea arenaria (Choisy) Steud. Nom. 19. Bot., ed. 2. 815. 1841, non Roem. & Schult. Ipomoea egge rsiana Peter in Engler & DT Nat. Pflanzenfam. IV (3a): 30. 1891, nom. d. Distribution: Puerto Rico, St. Croix, Virgin Gorda, St. Thomas, St. John. Choisy based Exogonium arenarium on four collections. These collections came from Puerto Rico, St. Thomas, Santo Domingo, and the Bahamas. Accord- ing to the interpretation that has been used for the past 60 to 70 years, the species does not occur on either the island of Hispaniola or in the Bahamas. Therefore, neither of the collections cited by Choisy from these islands should be chosen as the type. The specimen in Geneva matches the concept of historic use and has been chosen to be the lectotype. 22. Ipomoea avicola D. Austin, nom. nov. Basionym: Exogonium verruculosum Pittier, J. Wash. Acad. Sci. 21: 142. 1931. type: Venezuela, Aragua, Pittier 12118 (VEN, holotype; G, US, NY, isotypes ). Synonym: Ipomoea verruculosa (Pittier) O'Donell, Lilloa 26: 379. 1953, non I. verru- culosa Mart. ex Choisy in DC., Prodr. 9: 378. 1845, nom. pro syn 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Distribution: Endemic to the coastal mountains of Venezuela. This and the Brasilian I. longistaminea O'Donell are similar. These two populations apparently represent local endemic bird-pollinated flowers derived independently. 23. Ipomoea viridiflora Urban, Symb. Antil. 3: 348. 1902. TYPE: Ehrenberg 345 (US, isotype). Synonym: Exogonium viridiflorum (Urban) House, Bull. Torrey Bot. Club 35: 106. 1908. Distribution: Cuba, Hispaniola. Group 7. These two species belong to different alliances within the Erio- spermum group of Ipomoea. Flowers on the plants are apparently bee pollinated and the inclusion of these species in Exogonium appears anomalous. I. Ipomoea argentea Meisn. in Mart., Fl. Bras. 7: 247. 1869. svNTYPES: Brasil, Gardner 3356 (not seen). Venezuela, Spruce 3605 (K Synonyms: Ipomoea comosa House, Ann. New York Acad. Sci. 18: 201. 1908. TYPE: Based on I. villosa ( Choisy) Meisn. Batatas villosa Choisy in DC., Prodr. 9: 337. 1845. TYPE: Brasil, Martius 609 (M, syntype). Ipomoea villosa (Choisy ) Meisn. in Mart., Fl. Bras. 4. 1869, non Ruiz & Pavon, i. Exogonium villosum Choisy) Peter in Engler & Prantl, Nat. Pflanzenfam. B (3a): 28. 1891. Distribution: Known from savannas in Venezuela, south to Brasil. The plants show considerable variation throughout the range, prompting the division of the population. O'Donell (1960) was one of the first to point out that they were the same. 2. Ipomoea steerei (Standley) L. O. Williams, Fieldiana, Bot. 32: 195. 1970. Basionym: Exogonium steerei Standley, Carnegie Inst. Wash. 461: 83. 1935. TYPE: Steere 1545 (F, lectotype). Steere 1599 (F, syntype). Distribution: Reported from Mexico and Guatemala. LITERATURE CITED Austin, D. F. 1975a. Typification of the New World subdivisions of Ipomoea. Taxon 24: 107-110. ; 5b. Sed be ae tia In R. E. Woodson & R. W. Schery, Flora of Panama. Ann. Missouri Bot. Gard. 62: 157-224 Cuorsy, J. D. 18: Co 1 Orientales. Mém. Soc. Phys. Genéve 6: 385-502 gee 25 Convolvulaceis Dissert: em . Mém. Soc. Phys. Genève 8: 121- 164, Bee oo: [9v] ere "laceae. In A. de Candolle, Prodromus Systematis Naturalis Regni 5 Vol. 9: 323-465. Farcri, K. & L. VAN DER PIII. 1971. The Principles of Pollination Ecology. Ed. 2. Per- gamon Press, New York. GniskBACH, A. H. R. 1864. Flora of the British West Indian Islands. Lovell Reeve & Co., London. House, H. D. 1908. Studies in the North 7 Convolvulaceae Exogonium. Bull. Torrey Bot. Club 35: 97-107, 7 LioGign, A. H. 1968. Novitates Antillanae. III. ern id 152. Marupa, E. 1963 (1964). El genero Ipomoea en Mexico—l. Anales Inst. Biol. Univ. 5. IV. The Genus 1964 (1965). El genero Ipomoea en Mexico (II). Anales Inst. Biol. Univ. Nac. México 35: 45-76. 1977] AUSTIN—REALIGNMENT OF EXOGONIUM 339 65 (1966). El genero Ipomoea en Mexico (III). Anales Inst. Biol. Univ. Nac. 106. Meeuse, B. J. D. 1961. The Story of Pollination. Ronald Press, New York. MEISNER, C. F 3 Convolvulaceae. In K. F. P. von Martius, Flora Brasiliensis. Vol. : 20 O’DoNELL, C. A. 959. Las especies americanas de Ipomoea L. sect Quamoclit (Moench) Griseb. Lilloa 29: 19-86. 1960. Notas sobre Convolvulaceas americanas. e 30: 39-69. PERC ns M.S. 1965. Floral Biology. Pergamon Press, Lon PIJL, VAN DER. 1960. Ecological aspects of flower evalifiss I. Phyletic evolution. a 14: 416—463. 19 Ecological aspects of flower evolution II. Zoophilous flower classes. Evo- lution 15: 44— 59. ROhERTSON, K. R. 1971. A revision of the genus Jacquemontia (Convolvulaceae) in North and Central America and the West Indies. Ph.D. dissertation, Washington University, St. Louis. 285 pp. SrANDLEY, P. C. & L. O. WILLIAMS. 1970. Convolvulaceae. In Flora of Guatemala. Fieldiana, Bot. 24(9): 4-85 VERDCOURT, B. 1957. Typification of the ee of Ipomoea L. (Convolvulaceae ) with particular regard to the East African species. Taxon 6: 150-152. Convolvulaceae. In C. E. Hubb: 110 & E. Milne-Redhead (editors), Flora of Tropical East Africa. Crown Agents for Oversea Governments and Administrations, London. THE LUPINUS MONTANUS COMPLEX OF MEXICO AND CENTRAL AMERICA Davip B. DuNN? AND WILLIAM E. HARMON? ABSTRACT The recognition of the Lupinus montanus complex by morphological traits is discussed. Ecological modification of traits is discussed and the island nature of distribution from moun- tain peak to mountain peak produces semi-isolated gene pools. Long range dispersal and intro- zu s Potosí, Mexico, developing L. cacuminus. A similar situation occurred in Costa Rica, with L. valerioi the product of introgression from, as yet, a J m, n unknown t austrovolcanicus represents local introgression from L. kellermanianus, into L. montanus. Both of the Peruvian (L. praestabilis and L. proculaustrinus) taxa are, likewise the result of long range dispersal and introgression. The geographic range of each of the taxa of the complex is plotted and the interrelationship is discussed. The alkaloids have been plotted from random samples of each of the taxa and the data supports the taxonomic treatment and interpretation of their interrelationship. The lupines of Mexico have never been studied monographically. Previous studies have been floristic for states or regions or miscellaneous descriptions, as contributions to the flora of Mexico. To avoid further nomenclatural complica- tions, the earliest named taxa should be identified first. In this sense, the first taxon named for Mexico was Lupinus mexicanus Cerv. ex. Lag. (1816), which has been identified (Dunn, 1972). Lupinus montanus H.B.K. (1823) was the second epithet published for Mexico, concurrently with L. elegans H.B.K. (Humboldt et al., 1823: 478). Both of the types of these taxa are available at Paris, France, with microfiche illustrations now widely distributed. The topo- type material was studied and dissections of 50 collections, representing the geographic range of L. montanus were made, and the mean measurements were used to prepare the illustrations of L. montanus and allies presented in this paper. The illustration of L. montanus was sent to Paris and the curator of the herbarium kindly varified that the illustration accurately represents the species by matching it with the type specimen. Since L. elegans H.B.K. is the first epithet in a different complex of lupines, it will be treated, as soon as the rest of the complex is understood. With this approach it is believed, after ten years of study of the Mexican lupines and dissection of over 100 types for Mexico, that the taxonomic treatment of the L. montanus complex for Mexico and Central America can be presented. C. P. Smith (1948: 608) reported two South Ameri- The authors wish to express appreciation to the multiple curators of herbaria who loaned material for the study, as cited in the distributions by the code letters from Index Herbariorum Holmgren & Keuken, 1974). Special appreciation is expressed to the curators at Paris for comparing the illustration prepared for L. montanus with the type specimen. Two additional de. C herbaria are cited which were not in the code. — University of Northern Colorado, Greeley, Colorado, U.S.A., and WUP = Wisconsin University—Platteville, Platteville, Wiscon- sin, U.S ? Division of Biological Sciences, University of Missouri-Columbia, Columbia, Missouri 65201. ? Department of Biology, University of Northern Colorado, Greeley, Colorado 80631. ANN. Missouni. Bor. Garb. 64: 340-365. 1977. 1977] DUNN & HARMON—LUPINUS 341 can species to belong to what he interpreted as the “Lupinus montanus com- plex.” Only one of these was available for study. It is a member of the complex and is very distinctive in multiple characteristics. Both are only treated in the key. MORPHOLOGICAL RECOGNITION Lupinus montanus, per se, can be readily identified by the large sheathing stipules, the largest per specimen being 3-33 cm long. The infraspecific taxa show various modifications of the shape, size, texture, and vestiture of the stip- ules. (The allies are so different that they cannot be recognized by the stipules.) The stems, 0.5-2.5 m tall (to 4 m shrubs in Peru), are clustered from a woody caudex, and in age become short woody trunks, to 5 cm diameter. The current years growth is normally hollow and fistulose or subfistulose, varying from 5- 30 mm in diameter. (Allies and products of introgression have woody, solic stems, as small as 3 mm in diameter, and are only 3-7 dm tall.) The leaves show no general relationship other than a tendency to have many (9-17) leaf- lets, palmately compound, linear-elliptic to oblanceolate and generally glabrous above, but the last trait is modified by increasing hairiness above, with higher elevations and by introgression from the allies. The flower structure shows the greatest degree of uniformity, with only very subtle changes in size, shape and vestiture of the calyx, and size and position of the bracteoles. Keels are gen- erally glabrous but both infraspecific taxa and the allies may have ciliate keels. The base of the deep sulcus of the banner appears to have a nectary, which is uncommon in Lupinus. The number of ovules varies from taxon to taxon, with L. montanus and the infraspecific taxa varying from 7-13 per pod (the allies less). The seeds of L. montanus and the infraspecific taxa are very similar in shape to those of L. polyphyllus, a species distributed from southern California to British Columbia, with which it has been confused. The seeds are generally 4.5 mm long and 3 mm wide, with a deep funicular pit at the side of one end. While the large stipules represent the most distinctive trait of the L. mon- tanus complex, they are very small in some of the allies and reduced in taxa considered to be products of introgression. The large flowers with the banners reflexing near the midpoint are perhaps the most consistent character of the complex. They may indicate the utilization of specific pollinators. HABITAT AND ECOLOGICAL MODIFICATIONS The taxa of the complex are associated with the upper forest openings extend- ing upward to timberline. Thus the population on each mountain represents a breeding population with a chance for some genetic drift or fixation. This is reflected by variations in the hair type from mountain to mountain. However, to view the population on each mountain as having achieved some taxonomic status would be an exaggeration. It appears probable that migratory birds con- tribute to the separated populations of cach mountain peak intermittently. There is also an altitudinal modification of the density of pubescence of both the stems and leaflets. On Popocatepetl the smallest specimens were at the lowest altitude 342 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 and the area above timberline. The largest specimens were within the upper margins of timber. The densest pubescence was at the highest elevation and the sparsest pubescence was at the lower elevation, with the leaflets glabrous above at the lower elevations. On Nevada de Toluca, the type locality, the situation was similar. Sharma (1967, in an unpublished part of his thesis) demonstrated experimentally that hair frequency increased with aridity and heat. The vege- tative stature of the plants within the complex ranges from 0.3-2.5 m in height, while those within L. montanus vary from 0.5-2.5 m tall. EVOLUTION AND GEOGRAPHICAL RELATIONSHIPS (Fic. 1) Lupinus muelleri and L. kellermanianus, treated here as allies, can be uti- lized to illustrate both the islandlike mountain isolation and long range disjunc- tion of taxa with morphological similarities. The two are very closely related, but L. kellermanianus is known only from two volcanic peaks in Guatemala, while L. muelleri is known only from the opposite end of the distributional range of the complex on Cerro Potosí, Nuevo León, Mexico. Vegetatively they are very different from the rest of the L. montanus complex, having woody stems, low (3-5 dm) stature, only 2-2.5 cm long petioles, only 2-2.5 cm long leaflets with strigose-sericeous pubescence on both sides, and only 8-20 mm long stipules. The flowers, however, are very similar to L. montanus except that both are pubescent on the back of the banner. The vegetative traits, except pubescence, resemble L. argenteus of the Rocky Mountains of the United States but the closest geographic approach of this species is in northern New Mexico, while L. muelleri occurs in southern Nuevo León and disjunctly in Guatemala, where L. kellermanianus also occurs. The pubescence of the leaflets and the banner resembles that of the L. sericeus complex, but the closest approach of this complex is in northern Arizona, north of the Grand Canyon. If the source of the characteristics is from the above two complexes of the United States, then long range dispersal seems the only plausible explanation, via migratory seed- eating birds. Lupinus muelleri is on a mountain ridge surrounded by Chihua- huan desert, but inhabits the lower pine zone. Lupinus kellermanianus was collected high on Volcán Agua near timberline, hence both are in somewhat xeric situations where abundant pubescence is adaptive. INTROGRESSIVE HYBRIDIZATION Lupinus cacuminus is vegetatively intermediate between L. muelleri, with which it has geographic proximity, and L. montanus, from which it is geograph- ically completely isolated. Lupinus cacuminus has intermediate stipules, leaf- lets, and petioles, the fistulose stems of L. montanus, but the pubescent banner of L. muelleri, and extends above timberline, an ecological trait of L. montanus. In some populations of L. cacuminus an intermediate amount of ciliation occurs on the keel, a trait derived from L. muelleri. The multiple collections are very similar and appear to be a stabilized entity, derived from the hybridization of the L. montanus genome with the L. muelleri genome. The alkaloid spectrum of L. cacuminus is identical with that of L. montanus as shown below. 1977] DUNN & HARMON—LUPINUS 343 @=Lupinus montanus = r. nelsonii QzL. m. var. austrovolcanicus tesii ssp. glabrior +=Lupinus cacuminis W-Lupinus muelleri O=Lupinus kellermanianus X=Lupinus valerioi "IURE l. Distribution of the taxa of the Lupinus montanus complex and their allies in Mexico Ah Central America. Lupinus montanus subsp. montanus var. austrovolcanicus may be the prod- uct of introgression between ancestral L. montanus and L. kellermanianus. In this case L. kellermanianus could have provided the woody stem, reduced stat- ure, and intermediate vegetative traits. Both L. kellermanianus and L. mon- tanus var. austrovolcanicus are limited to one or two mountain peaks and have only been collected a few times. It is thus questionable whether the process of introgression has progressed long enough to have established a stabilized taxon of intermediate appearance. Another possible example of a long range introduction of some portion of the L. montanus genome into a lupine population is provided by L. valerioi on Cerro Vueltas in Costa Rica. There is a long distance between this population and the nearest known material of L. montanus in Guatemala. In this case the stipules are intermediate in size, somewhat similar to those of L. cacuminus. The bracts, however, are quite large and broad, typical of L. montanus. The stems are very slender, similar to those of L. kellermanianus, but the petioles are long, 6-12 cm, and have spreading pilose hairs to 4 mm long. These traits all suggest a mixing of genetic traits of L. montanus with some, as yet undeter- mined taxon of Lupinus. None presently known from Costa Rica provides these characteristics. A fourth example of long range introduction of genetic material from the 344 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 L. montanus genome is provided by L. proculaustrinus C. P. Smith of Peru. In this case the plants have retained many of the traits of L. montanus, including the large sheathing stipules and large broad bracts. The species has multiple unique traits, however, which are not present in L. montanus. A conspicuous one is the glabrous, glaucous surface and also a shrubby stature, reported on some specimens to be up to 4 meters in height. The other Peruvian species, L. praestabilis C. P. Smith has not been available for study. ALKALOID CHEMISTRY The material utilized was dried leaflets in all cases, since we have observed some cases where the alkaloids stored in the seeds were different from those present in the leaves. Seed material is not always present. Four to five leaflets were fragmented in a new coin envelope for each sample and transferred to a test tube. In species with large leaflets only one is necessary. Enough 30% KOH was added to wet the leaf fragments. Enough chloroform was added to cover the fragments. The rack with the test tubes was then stored in a refrig- erator for one day (24 h). A micropipette was utilized to spot 50 pl of the clear chloroform solution from the bottom of the test tube, for each sample, onto a thin-layer chromatographic plate (TLC). If no clear bottom layer was present, a few drops of chloroform was added to the test tube. The solvent utilized to separate the alkaloids was 95 parts chloroform, 4 parts anhydrous methyl alcohol, and 1 part ammonium hydroxide, by capillary flow, against gravity, in a Brinkman tank. The flow was stopped at 15 cm and the plate dried and developed first with Dragendorfs reagent. All visible spots were marked with a pencil. The plate was then sprayed with iodaplatinate to bring out any trace substances not observed with the first stain. The R, values plotted in Table 1 have been correlated with standards supplied by Cho & Martin (1971) for sparteine, lupanine, hydroxylupanine and cytisene, on each plate that was prepared. While the number of samples plotted is not large, it is quite clear that while some variation occurs in the trace alkaloids, the presence of the principal alka- loids is fairly consistent. Within L. montanus both varieties retain the same principal alkaloids, while the two subspecies show distinct alterations in the principal alkaloids. The suggestion that L. kellermanianus, L. valerioi, and L. cacuminus are allied is supported by the fact that the samples analyzed show the same principal alkaloids as those in L. montanus proper. Lupinus muelleri has a distinctly different spectrum of alkaloids from L. cacuminus suggesting that the two taxa are maintaining their isolation at the present time, even though they are in close proximity and are separated only altitudinally by a few hundred feet. The lone specimen available from Volcán Colima, probably in Jalisco, sug- gests that this population may be sufficiently isolated to require recognition. However, a single sample is not sufficient to permit an analysis of the situation, particularly since it appears to be morphologically very little different from the main population of L. montanus, even though it is geographically isolated. 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Jo ə5uəsəid əy} 10j po1so1 susundeds snuidn'] :junourP 9991] = L SS Əy} ozrpiep I 318v[Lp 346 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TAXONOMY Key ro Lupinus montanus AND ALLIES a. Banner sparsely strigose dorsally, near the distal half, along the crest; stipules 4.5 cm pe or less ee also var. V leaflets linear to linear-lanceolate, 2.5-5 cm long, ose to sericeous ab b. Plants mg 14 dm tall à or more, p anata or minutely 1 bracts per known only from Peru t treated) L. praestabilis bb. Plant 4-9 dm tall, sericeous to strigose; known from G MEA a or Mex etioles 5-13 cm long; stems hollow, fistulose below; pubescence pes sericeous; Nuevo León, Mexico wk Mm 2. L. cacuminus cc Petioles 24.5 cm long; stems solid and ligneous d. Keels 3 pubescence of leaflets 5 sericeous; 1 fron Guatemala Pr PHONE 3 3 dd. ae pn pubescence of leaflets ‘strigose-se sericeous; aaa rom a and N » León ç aa. Banner glabr igiene stipules more than 4.5 cm long or the stems with spreading p be hairs, 3-4 mi P i leaflets mostly glabrous above, sometime 1 gose mild variable in size nae cm long. e. Pubescence of the stems ane abundant spreading pilose hairs to 4 mm long; stems becoming ligneous; petioles 6-12 cm long; largest leaflets 4-7 mm long; known from Costa Rica Lo oaleríoi ee. Pubescence of the stems strigose, glabrous, or canescent; petioles mubi in s stems ligneous or herbaceous; largest leaflets 5-15 cm long; widely ditsibured. f. Stems and stipules glabrous to minutely 1 leaflets glabrous above; keels glabrous or sparsely ciliate eot distally; known from Chihuahua, mar o, or Peru. Stems 1 19 0 wee glaucous; flowers 18-20 mm ongi shrubs to 3.5 cnown only from Peru not treated) L. proculaustrinus gg. Stems agree or ES rate; flowers 15-18 mm long; 3 stems 5 tall; known from northwestern Mexico. h. Boh lance dels strigose dorsally; upper lip of the calyx trun- cate with an irregular notch; pees from Bises ior and north- ern Durango CS ld. montanus sub osp. glabrior hh. inns lanceolate, the tips “attenuate ra setaceous the wer dorsal area glabrous; upper lip of the UE triangular; cnown from southwest Durango and Sinaloa — TT montanus subsp. montesii Stems, stipules, and bracts abundantly 5 1 je other are of Mexico and Guatemala. i. Stipules hispidulous to canescent within, the larget 10-33 cm long; stems fistulose, hispidulous to 5 p 12-33 mm in diameter; known only from Ixtlán, Oaxa exico . sie ete TUN — lb. L. 5 E montanus var. nelsonii ii. Rer glabrous ‘within, or if pubescent, not with the above combina- n of characteristics, generally less than 10 cm long; stems puberu- — > bae = = = — — > FIGURE 2. Tq aay of the typical structures of Lupinus montanus subsp. montanus var. montanus. The and vegetative parts are drawn e mean va a se 30 dissections from. ih. pem graphice range of the taxon. The lettering is the same on all of the go = i = lateral view of the left side of the flower; B = banner petal flattened, dorsal vie Sate outside lower portion, inside upper portion; Ca = calyx, cut at the left PART sinus and opened so that the insic 15 is illustrated; K = i n 5 enclosing staminal tube and pistil, with the mean number of ovules drawn; L = ge largest leaflet drawn to % the scale used for the floral part, the lower half gus "different hair types observed on the lower surface, the upper half shows hair types observed on the upper sur- — stem structure of first year growth and hair types, half scale; St — sheathing pair of stipules, half scale, showing hair types; W — wing petal. DUNN & HARMON—LUPINUS 347 7 s= >= = SS 348 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 lent to strigose or hispidulous, hollow, 4-12 mm in diameter on the first year's growth, becoming ligneous. j. Stems to 4 mm in diameter, but hollow on the first year's growth, finely appressed puberulent; ATE only 2—4.5 cm long an pubescent within; racemes genera = than 8 cm long; rare, known from ien án Santa Maria, Cua . L. montanus 1 5 montanus - var, . austrovolcanicus Stems of the re of cee growth over 5 mm in diame and flattening on pressing, variously eae m feris 4-9 cm long, ee glabrous over most of the area within; racemes over 15 cm long at maturity; known from Guatemala to central Mexico _....... la. L. montanus subsp. montanus var. montanus — =: la. Lupinus montanus H.B.K. subsp. montanus var. montanus, Nov. Gen. Sp. Pl. 6:477. 1823. rype: Mexico, Montosis Novae Hispaniae (Nevada de Toluca, 9,000-10,000 ft) (P, holotype, not seen; microfiche, MO).—Fic. 2. L. vaginatus Cham. & Schlecht., Linnaea 5:590. 1830. rype: Mexico, Monte Orizaba, Sep., Schiede & Deppe (HAL, holo otype; photos, NY, TEX, L. flabellaris Bertol., Fl. Guat. 30. 1840. TYPE: Guatemala, Volcán d'Agua (not seen). TOPO- ryPE: Harmon 3646 (MO, NY, UC, UMO ). Plants perennial, 0.8-2 m tall, rarely to P m; stems hollow above the ground level, with the current year's growth 5-12 mm in diameter, pubescence varying with locality and altitude, finely appressed puberulent, to strigose or sericeous, more densely at higher altitudes, or hispidulous to retrorsely hispidulous or canescent; petioles of the primary leaves along the upper stems 6-15 cm long with the stipules connate nearly half the length of the petioles, those of dwarf lateral branches often reduced in size; stipules ensheathing % or more of the diameter of the stems, 4-9 cm long, the triangular free tips 4-12 mm long, both petioles and stipules pubescent dorsally as the stems, the stipules generally glabrous or glabrate within, on dwarf branches the stipules of the multiple leaves imbricated; leaflets of the larger primary leaves 10-15, linear to narrowly oblanceolate, the largest 5-13 cm long, 5-14 mm wide, generally glabrous above at lower elevations and puberulent to strigose above at higher elevations, the tips acute or slightly attenuate; peduncles 10-22 cm long, shorter on late-season branches; racemes 15-30 cm long at maturity, verticillate to subverticillate; bracts large-sheathing or covering 3-5 cm of the tip of the elongating raceme, hiding the buds, the tips attenuate-caudate, pubescence as the stipules for each population, generally caducous; pedicels 5.5-8.4 mm long, hispid or with appressed hairs; calyces sericeous, strigose or canescent on the outside, puberu- lent within on the distal portion of both lips, the lower lip 8-11 mm long, gen- erally entire, the upper lip 5.5-8.4 mm long, the notch at the tip 1-5 mm deep, the lips connate laterally 1.4-1.8 mm, the bracteoles 1.0-3.4 mm long, attached below the lips of the lateral sinuses; corollas glabrous, blue to lavender or purple, occasionally white or pink; banner obovate-rotund, longer than wide, the tip emarginate, 12.6-14.8 mm long, 11-14.5 mm wide, reflexed near the midpoint, reflexed 5.7-8 mm, appressed 6.7-7.8 mm, reflexed/appressed ratio 0.81-1.06, the angle 130?-146^; wings 13.8-16 mm long, 6-10 mm wide, the claw 2-3.7 mm long; keel 4-5 mm wide in the middle, the angle 84°-98° (aver- age 90.9°); ovules 7-10; pods 4-5 cm long, 9-10 mm wide when dried, arching m 1977] DUNN & HARMON—LUPINUS 349 up and outward, abundantly to sparsely tangled-pilose, the hairs 1-2.5 mm long; seeds black or brown with dark mottling, 4-4.5 mm long, 3-3.6 mm wide, with a deep funicular pit; chromosome number n = 24 (Beaman et al., 1962). This is one of the wide-ranging species of the volcano zone of Mexico and Central America. It occupies a zone on most of the high mountains from tim- berline, or above in sheltered areas, down through the upper pine forest into the mixed pine-oak forests. The dominant alkaloid produced is sparteine, with only traces of minor alkaloids. Nowacki's (1963) contention that sparteine is the primitive alkaloid would suggest that this may be one of the older, more primitive species of Lupinus in North America. The trait of many leaflets and the fistulose stems have caused many botanists to mistake this taxon for L. polyphyllus which ranges from California and Oregon north into British Colum- bia. The huge sheathing stipules, however, make L. montanus easily recog- nizable. While L. polyphyllus is considered as one of the older taxa of the West Coast, it has the necessary genes to convert or utilize sparteine, changing it into several other alkaloids. The two northern subspecies of L. montanus have also modified genomes so that they concentrate other alkaloids. The presence of what appears to be a distinct nectary at the base of the ventral sulcus of the banner and the thickened glandlike upper surface of the base of the staminal tube seem to be unique in Lupinus. While the original description of L. montanus failed to mention the sheath- ing stipules, they are clearly recognizable in the microfiche of the type speci- men, and the material from Mt. Orizaba differs from that of Nevada de Toluca only in the hair type being hispidulous to canescent, hence the contention that L. vaginatus is a synonym. The type description of L. flabellaris clearly men- tions the sheathing stipules and topotype material is indistinguishable from the Mexican portion of the taxon, both morphologically and chromatographically. GUATEMALA. DEP. CHIMALTENANGO: Chinar, Skutch 107(US). Chichoy Pros, Hun- newell 17145(GH). San Marcos, I. R. Johnston 1229(F). Sierra San Elna, Siler 2303( GH). nes 7 ango, Beaman 3258(GH 1 8 TEX, US); Standley 61902 (F). Volcán de Agua, R. Johnston 578 (F). Volcán Fue N side, Beaman 4025(GH, MSC, US). DEP. HUEHUETENANGO: Cumbre Papal, Ste at 0% ). Between Tocquia and S San Juan 5 tm Ixcoy, Moncure in 1950 (F). Between Tunima and Quisil, Steyermark 48426(F). DEP. Q ZALTENANGO: Canton La Eseranza Forest, 6 km from San Juan Ostuncalo, Molina et al. 16648 (F, US). Cerro Lieteoreya, Koninck 302(U S). 10 mi W of Quezaltenango, King 3199 (MICH, TEX, US). 11 mi SE of San Marcos, King 3173( MICH, TEX, US). Sierra Madre Mts., Williams et al. 29780(F, US, WIS), 2 2800 ( F). Volcan Santa MED: Skutch 855( F, GH, US): Steyermark 34168(F). Volcán Santo Tomás, Steyermark 34857 (F). DEP. SACATEPE- GEZ: Volcán Agua, Harmon 3598( UMO, US, WIS), 3646( CAS, CUN, F, MICH, MO, UC, UMO, US); Beaman 2942(GH, MSC, TEX, US); Kellerman 4708( U S); Maxon & Hay 3687 (US); Molina 21029(F ); Salas in Jan. 1926(US); Shannon 3679(US); J. D. Smith 2152 (US); Standley 65093(F). pep. SAN marcos: Cerro El Bonito, Plowman 5044( GH). Be- tween San Sebastian and summit, Steyermark 35553(F ); — ie s ion 35530(F). 2 mi S of San Sebastian, Williams et al. 25915(F, WIS). Volcán Tacaná, Beaman 3230(GH, MSC, TEX, US); Steyermark 36140( V). Volcán HN dm ee ASH. MSC, TEX, US); Reeder in Apr. 1952( MICH ); Shannon 568( US); Williams et al. 26996( F, GH, US). DEP. SOLOLA: Volcán Atitlán, Beaman 4094(GH, MSC); Kellerman 5769(US); Steye rmark 47508(F, US). Volcán Santa Clara, Steyermark 4698S(F). Volcán Tolimán, Steyermark 47535(F). DEP. TOTONICAPAN: Cerro María Tecüm, Williams et al. 23145(F). Boundary of Depts. Huehuetenango and Quezaltenango, W illiams et al. 22707(F, GH, US, WIS). Los — — 350 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Encuentros y gate Tecùm, Molina 15879( F, US, WIS). 8 km S of Totonicapán, Williams et al. 22941 (F, US); Lind 11(V, WIS MEXICO. CHIAPAS: Mt. Pasitár, Matuda 70(MICH, NY, US); Matuda US MICHI. Volcán Tacaná, 2d 2333(GH, MICH, WIS). corma: Cuchilla, Nevada de Colima, d 57(F , MO, US). DISTRITO FEDERAL: Cerro Ajusco, Beaman 9796(CH, MSC, US); Garcia in ue 1954(IPN). Pedregal de San Angel, Barclay & Paxton 530(F, TEX). de de los Charros, Russell & Souviron 139( US). GUERRERO: Mina, Cerro Teotepec, Hinton 4266(GH, MICH, US); Rzedowski 16493( ENCB, MICH). nipArco: Plains of Actopan, Co 168 H). Cerro de las Venturas, N of Pachuca, Galvan in Aug. 1963 (ENCB); Nunez 56(UMO). El Chico, Dunn et al. 20298(UMO, US). Sierra de Pachuca, Rose & Hay 5628( US); dod 9530(F, GH, MO, US, VT). janisco: Nevada de Colima, Beaman 2366 (MICH, MSC, US); Matuda 38369( MEX, UMO); McVaugh 10076( MICH, small flowers). MEXICO: 10 km E of Amecameca, Quijana 51(ENCB). 11 mi E of Amemeca, Dodds 11 (MICH); Montgomery & Root 8114a( MSC). Cerro Jocotitlán, Matuda 38490( MEX, UMO). CruceroPAgua-Blanca, Hinton 8244( GH), 8317( GH, MO, RSA, UMO, US). Crucero-Raices, Hinton 9031(F, GH, ENCB, MO, RSA, UMO, US). Estacca, Matuda 38503( MEX, UMO). Ixtachuatl, Falda, Matuda 26148(NY, US); Beaman 3477(GH, MSC, US, n — 24); Nelson & Goldman in Jan. 1894( US); Purpus 32( US), 208( MO); Rudd 1029( UMO, US); Rzedowski 19806( ENCE, UMO). Mesón Viejo, Matuda 38395( MEX, UMO. Nevada de Toluca, Balls ipie et al 19 915(M EMO, MO, NY, UC, UMO, WUP), 1932( ASU, GH, MO, MSC, OSC, ) UMO), 1936( MEMO, MO, RSA, UMO, US, WUP); Gallian & Leake 897 EMO Galliotti 3360(P); Hunnewell 13146( GH); Islas 28(MEMO); Mick & Roe 187 (ENCB); Morales-Diaz in Aug. 1962( ENCB); Rose & Painter 7906(US); Rzedowski 15782 (ENCB); Schery 90( MICH, MO, US). Paso de Cortez, Iltis et al. 1025 (MICH, MSC, TEX, WIS). Pesco Inst. de Nacional, Dunn et al. 20373(F, GA, K, ENCB, MEMO, MO, NEL, NY, ORE, RSA, UC, UMO, US, WUP). Río Frio, Mii dn ^ July 1962( ENCB). Tlamacas, vic. of Popocatepetl, Fonseca F5(ENCB); Galicia in July 1962( ENCB); Garcia 3A( ENCB); Lundell 12358( MICH, TEX); Madrigal in Dec. 1959( E. Matuda 38497 (MEX, UMO); Moore 36(GH); Quijano 51(ENCB). Volcan Popocatepetl, Balls 4227( US); Barkley et al. 1 0 Y; Beaman 2022(MSC); Dunn 18566(UMO); 185790 CUN, ENCB, MO, NY, UMO, US); Galliotti 3368(P); Hatheway 1193( GH, MO, US); Huerta 101( ENCB); Leake & Gallian 133, I41( UMO); Lundell 12358( US ); Rose & Hay reci Ross S(US); Straw & Gregory 1003(GH, MICH, RSA). micHoacAN: Cerro San Andres, 12 km N of Hidalgo, Beaman 4295(GH, MSC, TEX, US). Mt. Tancitaro, Hinton ae Leavenworth & Hoogstraal 1128(F, MO). Zitácuaro-Cacique Peak, Hinkson 11932(US). oaxaca: Atepec, Llano de las Flores, Mac Dousal 378.S(NY). Cerro de San Felipe, Camp 2869( NY). Cerro 5 Hallberg 790(ENCB, MICH, US). Cumbre de Sierra de Juárez, — 38415( MEX, UMO). Gueletago, Vilas 31( WIS). 27 mi N of Ixtlán, Sierra Juárez, Roe Roe 1941 (ENCB, MICH, WIS). Macuiltianguis, MacDougal in 1960(US). Mt. 5 summit, E. Nelson 619008). Reyes, E. Nelson 1736( MICH » US). Sierra de Ixtlan, Gentry et al. 20272(UMO). Sierra Madre del Sur, 60 mi NE of Oaxaca, Webster 11543( MO). Sierra de San Felipe, Camp 2869( NY); E. Nelson 1135(US); Pringle 4779(F, GH, ISC, MICH, SC, ND-C, P, US, VT); C. L. Smith 333(MO). ruEBLa: Iztaccihuatl, ` Beaman 2007 ( GH MSC, US); Iltis et al. 1025(TEX); Weber 372(ENCB). Pass between indeed "yd and Puebla, Mexia 2647 (MICH), 2647 A(CAS). Alberque 3 Grande, Beaman 364 MSC, TEX. Ciudad Serdan, Beaman 2498(GH, MSC). Pico de Orizaba, 10 Sa Greenman 28(F); Liebman 4892(F, GH), 4893( F); Panels 9528(F, O, US, VT); Schiede 666( HAL; photos, GH, TEX, UMO, US); Eee on 510(GH). PE Barkley 17Mo87(F); Barkley et al. 2353; Beaman 1747, 2109(GH, MSC); Dunn 18558(UMO), 18564( CUN, ENCB, MO, NY, UMO, US); 7 7 e & Barkley 17MoS7 (TEX, MSC). TLAX- CALA: Mt. Malinche, Balls 4890 (US). veracruz: Cerro de Perote, Balls 4604( US). Cueva — > Ficure 3. Structures of Lupinus montanus subsp. montanus var. nelsonii drawn to mean values, for those traits which differ from var. montanus. The calyx, bract and stem are drawn to the scale shown, while the ir pue nd leaflet are half ae The lettering is Br = bract; Ca = calyx, inside view; L = caflet: S — stem; St — stipules. (See legend for Fig. 2 for full explanation.) 351 1977] DUNN & HARMON—LUPINUS ~ 10to33CM— St! . " E e" % » z aS " p ` `. 27 e ` ^ 5; en ta y af ` * " ` . NV Fam p na NIS z ' q £ yf k 1 oo ot ` £ S. x \ ` Fa uo ` roy LA k: / \ ' ` i E 22 ' , t i ` «t `. \ É a va 1 `. b 1 + LAS ^ HE x ç ` ' k E ' 8 am t1 * ii 14 ' Ti va ` 1 Ei Ë s * + 1 1 ' L * 3 1 5 vi fo ` ' ' ' ` & oq ti ti T Fog ' moa 1 i3 '. ' ! roe Ñ . ck "n Ë: ' ' ' 1 E ` ' C e ` ` yx D ` B e q v var. nelsonii (Rose) C.P. Sm. Lupinus montanus H.B.K. 359 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 de Muerto, Beaman 1783( MSC, US). Pico de Orizaba, Beaman 2291(MSC); E. Nelson 266 US); Rose & Hay 5727(US); T G. Smith 391(MO). Unknown locality, E. Nelson 38( US). 1b. Lupinus montanus subsp. montanus var. nelsonii (Rose) C. P. Smith, Sp. Lup. 79. 1938.—Fic. 3 L. nelsonii Rose, Contr. U.S. Natl. Herb. 8:308. 1905. type: Mexico, Oaxaca, near Cerro de san Felipe, E. W. Nelson 1145(US, holotype, photo, UMO). Differs from L. montanus in the large fistulose stems, up to 3 cm or more in diameter, the fistulose nature extending throughout the above ground por- tion to the top of the raceme; plants to 2.5 m tall; stems densely hispidulous to retrorsely hispidulous; petioles to 50 cm long; leaflets 15, to 15 cm long and 3 cm wide, pilose to canescent below and glabrous above; stipules 10-33 cm long, connate to the petioles for all but 1-2 cm at the tip, the free tips slender attenuate-caudate, both stipules and bracts hispidulous-canescent within as well as densely so without; bracts numerous, plumed, the tips elongate-caudate, 3— 4.5 cm long, pilose-canescent, as also the under side of the leaflets, the bracts hiding 5-8 cm of the buds at the tips of the racemes, often subpersistent; flow- ers the same as the species; pods and seeds the same as the species except the pods densely hispidulous. The taxon appears to be a gigas form and may represent an ecological mod- ification since there are many typical specimens of L. montanus in the region. There are distinctive traits, however, which appear to have a genetic basis, and the taxon has been collected on several occasions from 1894 to 1964. It is also chromatographically similar in its alkaloids to other samples of L. montanus. Mexico. OAXACA: Ixtlan, Atepec, Llanos de las Flores, MacDougal 378s( NY). Oaxaca- Tuxtepec Hwy., Llanos de las Flores, MacDougal in 1960 (US, 4 sheets). Cerro de San Felipe, E. Nelson 1145(US); Pringle 5839(GH, VT). Cerro Zemoaltepetl, Schultes 502(GH); Hallberg 790( MICH, in part). Ixtlán de Juárez, Sierra Ixtlan, Gentry 20272( UMO, US). 13 mi N of Ixtlan, Anderson 4842( MICH). lc. Lupinus montanus subsp. montanus var. austrovolcanicus C. P. Smith, Sp. Lup. 90. 1938. type: Guatemala, Volcán Santa María, 8,000-11,500 ít, E. Nelson 3709( US-250873, holotype; F, GH, isotypes ).—Fic. 8. Plants perennial, over 3 dm tall; stems hollow, ligneous, 4 mm in diameter, finely appressed puberulent; petioles 6-10 cm long; larger stipules 3-4.5 cm long, wide, membranous, sparsely pilose to canescent inside and outside, the free tips 7-11 mm long; leaflets 10-11, linear, acute, mucronate, the longest 5.5-6.5 cm long, 6-8 mm wide, sparsely strigose above, thinly kinky canescent beneath; peduncles 2-8 cm long; racemes ca. 8 cm long, verticillate, the lower whorls to 2 cm distant; bracts ca. 16 mm long below; pedicels 5-6 mm long, slender sericeous-puberulent; calyces canescent without, finely sericeous within near the tips of the lips, the lower lip slender, arcuate, 8-9 mm long, entire, the upper lip ovate, bidentate, 7 mm long, the lips connate laterally 2 mm, the bracteoles 1.5-2 mm long, attached near the lip of the lateral sinuses; banner suborbicular, glabrous or occasionally sparsely hairy on the distal portion of the dorsal crest, 11.5-12 mm long, 11-11.5 mm wide, widest above the midpoint, reflexed 6.7 mm, appressed 7 mm; wings 12.8-13 mm long, 7 mm wide; keel 1977] DUNN & HARMON—LUPINUS 353 with minute papillae above the claws, 3.5 mm wide in the middle, the angle 5°-100° at anthesis; ovules 6-7; pods 34.5 cm long, 9 mm wide, pilose with hairs 1-2 mm long. The specimens seen appear to represent hybridization and introgression from L. kellermannii. The slender woody stems, short stature of the plants, narrow smaller leaflets, intermediate petioles, and the presence of pubescence dorsally on the banners of about half of the specimens all suggest introgression. The flower size and stipules are distinctly like those of L. montanus. GUATEMALA. DEP. QUEZALTENANGO: Volcán Santa Maria, Beaman 4124(ENCB, M F an ASC, , US); E. Nelson 3709(F, GH, US); Steyermark 34205(F). Above Palojunoj, Standley 67703, 67707, 67738(F ), 67683(F, intermediate to var. montanus). 1d. Lupinus montanus subsp. glabrior (Wats.) Dunn & Harmon, comb. nov. —Fic. 4 L. montanus var. glabrior Wats., Proc. Amer. Acad. Arts 23:270. 1888. rype: Mexico, Chi- huahua, summit of Sierra Madre, Pringle 1206( GH, holotype; F, G, K, ND-G, P, RSA, T, isotypes; photo, UMO L. glabrior ( Wats.) Rose, Contr. U.S. Natl. Herb. 8:308. 1905. Plants perennial, to 1 m tall; stems glabrous to glabrate, ligneous ridged, at least on drying, fistulose, 6-8 mm in diameter; petioles 12-20 cm long, the free portion finely strigose; stipules membranous, sheathing and often imbricate on dwarfed branches, 4.5-6 cm long, the free tips only 3-6 mm long; leaflets 14- 15, the largest 9-11 cm long, 12-13 mm wide with acute-mucronate tips, gla- brous above, sparsely and finely puberulent below; peduncles to 17 cm long; racemes 20-30 cm long at maturity, verticillate, the lower whorls, 2.5-3 cm distant, the rachis finely but densely puberulent; bracts broadly lanceolate, membranous, gradated, the lower to 2 cm long, the upper reduced, minutely puberulent without, glabrous within; pedicels 6-8 mm long; calyces with broad boat-shaped lips, finely puberulent without, glabrous within, the lower lip 7-8 mm long, entire, the upper lip 5-7 mm long, the apex blunt with an irregular notch 0.5-0.8 mm deep, the lips connate 1.4 mm, the bracteoles straplike, 2-3 mm long, glabrous, except for a few setaceous hairs near the tips; corollas gla- brous except for a few papillae above near the claws of the keel or occasionally the keel ciliate above toward the acumen; banner orbicular, 13-14 mm long, 13-15 mm wide, reflexed 7 mm, appressed 6.5-6.8 mm, the sulcus 2.4 mm deep midway between the umbo and the base, the banner angle 133°-150°; wings 15-17 mm long, 8-10 mm wide; keel 4-5.3 mm wide in the middle, the angle 0°-85°, occasionally papillae near the claws or occasionally ciliate above near the acumen; ovules 7-9; pods 8-8.5 mm wide, 3.5-4.5 cm long, thinly strigose; seeds nearly black with mottling, 4.5 mm long, 3 mm wide, a pit at the funic- ular attachment. The subspecies glabrior is known only from northern Durango and Chihua- hua, from the summit of the Sierra Madre Occidental, in rather inaccessible areas. The area is north of that of subsp. montesii and subsp. montanus, as well as the fact that there are distinctive morphological traits in addition to distinct [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN 354 T ————— "a Ww. zem uouupH S uung (sioM) 4011q0)6 'dsqns »8 ` H snuojuow snuidn7 1977] DUNN & HARMON—LUPINUS 355 chromatographic differences. The best distinguishing traits are the short, blunt upper lip of the calyx and the lanceolate bracts. While other traits are distinct, they are not as easily recognized. Since the geography, ecology, morphology, and chromatography suggest a distinct gene pool, subspecific rank is suggested. The large stipules and the floral morphology clearly indicate the affinity to L. montanus. MEXICO. CHIHUAHUA: Vic. of Chuchuichupa, summit of Sierra Madre, cium 1206(GH, a ie K, ND-G, P, RSA, US), 1579(P). Cerro Mohinora, 10 mi S of Guadelupe y Calvo, w & Foreman 1943( ENCB, MICH). puranco: Sierra Madre, 30 Bis N of Guanacevi, E. ‘Nelson 4785( US) le. Lupinus montanus subsp. montesii (C. P. Smith) Dunn & Harmon, comb nov. L. montesii C. P. Smith, Sp. Lup. 41. 1938. type: Mexico, Sinaloa, Cerro de San Rafael, San Ignacio, Montes & Salazar 112(US, holotype ).—Fic. 5. Plants perennial, from a woody caudex, 4-7 dm tall; stems hollow, glabrous below, sparsely strigose on the peduncle, rachis of raceme, and petioles, ligne- ous ridged, at least on drying, 6-8 mm in diameter; longest petioles 10-19 cm long, strigose on the portion not fused to the stipules; stipules sheathing and encircling % of the stem, 4-10 cm long, the free, caudate tips 1-2 cm long, glabrous except for a few scattered setae near the tips; leaflets 9-14, linear- elliptic to narrowly elliptic, glabrous above, sparsely strigose below, the largest 6.5-9 cm long, 6-12 mm wide, acute-mucronate at the tips; peduncles hollow, 7-14 cm long, sparsely strigose; racemes verticillate to subverticillate, 20—35 cm long, rarely only 7 cm long in depauperate specimens, the rachis more densely strigose; bracts membranous, the long caudate tips with scattered seta- ceous hairs, caducous, 1.0-3.5 cm long, broad and completely covering the flower buds at the tip of the raceme, pedicels filamentous, 5-8.8 mm long, densely strigose; calyces appressed puberulent outside, glabrous within, the lower lip 8-12 mm long, generally entire, occasionally with a notch 0.1 mm deep, the upper lip 6-9.5 mm long, with an apical notch 0.3-1.6 mm deep, the base gibbous above, the lips connate 1.2-2 mm, a glabrous spatulate bracteole 1-4 mm long attached near the lip of the lateral sinuses; corollas blue and white, glabrous but the keel sometimes ciliate; banner 11.6-15.9 mm long, 12.5-17.4 mm wide, reflexed 6.4—7.7 mm, appressed 5.5-7.4 mm, reflexed/appressed ratio (average 1.17), banner angle 1307-149? (average 141°); wings 13-18.4 mm long, 8.4-10.4 mm wide, the claw on the average 3.2 mm long; keel 3.8-5.2 mm wide in the middle, the angle 89°-98° (average 91°), ciliate above near the acumen in half of the specimens, the others glabrous; ovules 10-13; only imma- ture legumes seen, these strigose. < 4. Structures of Lupinus montanus subsp. glabrior drawn to mean values, for the differentes from subsp. montanus. All parts are drawn to the scale shown except the yema and the leaflet which are drawn to half scale. The lettering is: B = banner, dorsal Br = 555 Ca = 5 1 view; F = flower, left side view; K = kee = leaflet; S = stem; St = stipule; W = wing. (See legend for Fig. 2 for full meee s ) [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN 356 uou0H * uunq ("WS q'29)!!səluou; “dsqns 'M'H'H SNuDjuoW snuidn] `< cece ` “s. s, +s 7 n Os t< e OK — 1977] DUNN & HARMON—LUPINUS 357 The subspecies is known only from southwestern Durango, Mexico, in the eneral area of El Salto, and from Cerro de San Rafael, Sinaloa. Most of the collections have been west of El Salto but some have been cited as 45 miles to the south. The elevations have been near 7,000 ft, which is well below the normal altitude for L. montanus. The distribution is also geographically dis- tinct from that of the species. Since the ecology, geography, morphology, and the composition of the alkaloids, are distinctive, it is suggested that there is a sufficiently distinct gene pool to recognize the taxon at the subspecific level. It is also distinct from subspecies glabrior in geography, morphology, and in chromatography. The large sheathing stipules and the floral morphology leave no doubt as to its affinity with L. montanus. TEXICO. DURANGO: Cerro Auehueto, S of Huachicheles, Maysvilles 7258( MICH). Be- tween Durango and Mazatlan, Pennington 242, 243( TEX). El Salto, p in Sep. 1961 (ENCB). 20 km S of El Salto, Gordon 41( MICH, x L. madrensis). mi SW of El Salto, Waterfall 15490( OKLA). 33 mi SW of El Salto, Wate rfall 127 da UMO). 3mi W of La Ciudad, LeDoux et al. 2024(C, CAS, G, ENCB, K, MEX, MO, NY, U, UC, UMO, US). 6 mi W of La Ciudad, Flyr 274(TEX). 13 mi E OF La Ciudad, LeDoux et al. 1996(ENCB, ME} O, NY, UMO, US). 8.4 mi W of La Ciudad, Reveal & Alwood 3507 (MARY, UMO); Breedlove 18877(CAS, UMO). 14.3 mi NE of La Ciudad, Pinkava et al. 9495( ASU, UMO). 54 mi N of Estación Coyotes, dox 'diove 18789( CAS, ENCB, UMO). Metates, N of Cueva, Pennell 18408 (GH, US). Road to Pueblo Nuevo, Maysvilles 7760( MICH, x L. madrensis). W of Pueblo Nuevo, Maysvilles 8071( MICI I, TEX). siNALOA: Cerro de San Rafael, San Ignacio, Montes & Salazar 112(US). ho Lupinus cacuminus Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 79. 1940. type: Mexico, Nuevo León, peak of Cerro Potosi, Municipio de Gale- ana, Mueller 2269 (F, holotype; GH, MO, isotypes) [Mueller 1257 (F) labeled type in Standley's handwriting].—Fic. 6. Plants perennial, caespitose, 3.5-6 dm tall; stems from a caudex, fistulose, the internodes between fully developed leaves only 1-3.5 em long, pubescence all appressed-sericeous but of multiple hair types and lengths, the upper 2 or 3 nodes with branches initiated by anthesis of the primary racemes; largest peti- oles 5.5-13 cm long, reduced progressively upward, persistent long after the leaflets drop; stipules gradated from 4.5 cm long below to 1.5 cm above, imbri- cated below, connate 3 cm below to only 7-8 mm above, the free tips subulate- caudate; leaflets 10-14, linear-elliptic, appressed silky villous on both sides, sparsely above and the central area often glabrous, the largest 3-4 cm long, 3-4 mm wide (to 6 mm wide in a population on Peña Nevada), the tips acute and mucronate; peduncles 4-5 cm long, exceeded by the foliage; racemes 10-13 cm long but numerous bracts in a terminal tuft suggest that they may get much longer, the flowers tightly and spirally arranged; bracts lance-subulate, tardily deciduous or semipersistent; pedicels 6.5-12 mm long, hispidulous; calyces silky < E 5. Structures of Lupinus montanus subsp. montesii drawn to mean values, for the differences from subsp. montanus. All parts are drawn to the scale shown except those for the stipules and leaflet, which are drawn at B scale. The lettering is: — banner, dorsal view; Br — bract; Ca — calyx, inside view; F — flower, left side view; K — eaflet; S — stem; St — stipule; W — wing. (See legend for Fig. 2 for full E. ) [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN 358 ‘puos siuiunopo snuidnq 1977] DUNN & HARMON— LUPINUS 359 white-villous with appressed hairs, the lower lip oblong-lanceolate, 8-10.2 mm long, tridentate or entire, the teeth 0.2-0.4 mm deep, the upper lip 7.7-10.4 mm long, bifid, the notch 1-4 mm deep, the lip oblong, 3.5-4 mm wide, flattened, the lips connate 1.4-2 mm, the bracteoles lanceolate, 1.5-6 mm long, attached at the lip of the lateral sinuses; banner sparsely pubescent dorsally on the distal portion, the tip emarginate, 14-17.5 mm long, 13-17.6 mm wide, reflexed 7-10 mm, appressed 6-8.5 mm, the angle 140°-155°, the sulcus 1.5-1.8 mm deep midway; wings 15-17.4 mm long, 7-9.5 mm wide, the claw 2.5-3 mm long; keel 4-5 mm wide in the middle, glabrous or ciliate above near the acumen, the angle 80°-90° at anthesis; ovules 5-7; pods 4-5 cm long, 9-11 mm wide, densely lanate with hairs 2 mm long. While collectors have reported the plants as abundant in the upper pine woods and above timber line on Cerro Potosi, they have only been collected on three mountain peaks: Los Alpes, Cerro Potosi, and Peña Nevada. Flower- ing occurs from June through July and as late as August. The taxon appear intermediate between L. montanus and L. muelleri in several characteristics but has essentially the same spectrum of alkaloids as L. montanus. The speci- mens from Peña Nevada have sparse ciliation near the acumen of the keel. Mexico. COAHUILA: Los Alpes, 40 mi E of Saltillo, Geni et al. 20059( UMO, US). NUEVO LEÓN: Cerro Potosí, Municipio de Galeana, Mueller 2269(F, GH, MICH, MO). Biol. Exp., U. of Ill., Schneider 958(F, GH, MICH, MO). Cerro Pofg top DEG mt., Beaman 2654 (GH, MSC, US); Gilbert 26, 29(TEX); Hinton 17038( MICH). Cerro Potosí, near micro- wave tower, Dunn et al. 202 203(F, K, MO, NY, RSA, UC, UMO, US); Dziekanowski et al. 1761 ( CH, ENCB, K, MEMO, MEX, MICH, MO, MSC, UMO, WUP); MacGregor et al. 314(KANU, UMO). 20 mi E of Galeana, Mueller 1257(F, GH, MICH, TEX). Peña Nevada, 26 mi NE of Dr. Arroyo, top of Picachio Onófre, Beaman 2690(GH , MSC, US). TAMAULIPAS: Pena Nevada, E and S 858 Stanford et al. 2597 (DAO, U, US). Summit of Pena Nevada, . Gillett 1237( MSC). 3. Lupinus muelleri Standley, Publ. Field Mus. Nat. Hist, Bot. Ser. 22: 80. . TYPE: Mexico, Nuevo León, Las Canoas, on Cerro Potosí, Municipio de Galeana, Mueller 2205 (F, holotype; CAS, GH, MICH, MO, TEX, iso- types ).—Fic. 7 Q Plants perennial; stems few to many from a woody caudex, woody with a solid pith, erect, 5-7 dm tall, 3 mm in diameter, branching from the upper nodes, thinly appressed strigose, with a cinereous undercoat of kinky hairs 0.2-0.4 mm long; petioles 1-2 cm long, filiform; stipules subulate to filiform, 6-8 mm long, connate to the petioles 2-4 mm; leaflets 6-8, the largest 2-2.5 cm long, elliptic- oblanceolate, the tip acute, mucronate, both surfaces densely strigose; peduncles 2.5-3.5 cm long; racemes 6-10 cm long, the flowers scattered to subverticillate; bracts subulate, 8-8.5 mm long, strigose outside; pedicels 7-11 mm long, with < Ficure 6. Structures of Lupinus cacuminus drawn to the means values on the scale shown. The stipules and leaflet are drawn to half scale. The lettering is: B — banner, dorsal view; Br — bract; Ca — calyx, inside view; F = flower, left side view = keel; L = leaflet; S = stem; St = stipule; W = wing. (See legend for Fig. 2 for full 5 ) 360 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 hispidulous hairs 0.4 mm long; calyces sericeous outside, 2-lipped, the lower lip arcuate, boat shaped, 7.5-9 mm long, 3-4 mm wide, entire, or with teeth 0.2 mm long, the upper lip ovate, 7-7.4 mm long 3.8-4.5 mm wide toward the base, the apical notch 0.1-0.8 mm deep, the lips connate 1.2-1.4 mm, the bracteoles 0.8-2.0 mm long, linear, attached below the lateral sinus lips; banner sparsely pubescent dorsally near the distal portion, suborbicular, widest below the midpoint, reflexed 7.5-8.2 mm, appressed 6-7 mm, the angle 138°-145°, the sulcus 1.5-1.8 mm deep, midway; wings glabrous, obovate, 13-15 mm long, 7-7.9 mm wide; keel ciliate above from the middle toward the acumen, 4-5 mm wide in the middle, the angle 83°-88°; ovules 6-8; pods 4-5 cm long, 10-12 mm wide, thinly subap- pressed kinky pilose, the hairs to 1 mm long; seeds 6.4 mm long, 5.7 mm wide, tan with faint mottling. Thus far known only from Coahuila and Cerro Potosi in Nuevo Leon, where it was reported as abundant in pine woods, at elevations near 3,000 m. Flow- ering occurs from June to August XICO. COAHUILA: Puerto de la Siberio, Arteaga, Marroquin 14(MEMO). NUEVO LEON: Cerro Potosi, Las Canoas, Municipio de Galeana, Mueller 2205(CAS, F, GH, MICH, MO, TEX). Above Ejido, Beaman 3311(F, GH, ENCB, MSC, TEX); Dunn et al. 20247( UMO); Dziekanowski et al. 1770(CAS, G, ENCB, MO, NY, ORE, P, RSA, UC, UMO, US), 1771 (GH, ENCB, MEMO, MEX, MO, MSC, NY, RSA, SLP, UC, UMO, US, WUP). 4. Lupinus kellermanianus C. P. Smith, Sp. Lup. 90. 1938. Type: Guatemala, Volcán Agua, 9,000 ft, Kellerman 4746 (US, holotype ).—Fic. 8. Plants perennial, shrubby; woody stems solid, the branches sometimes hollow, 3-6 mm in diameter, first year stems only 3 mm in diameter, strigose; petioles filiform, 2-4.5 cm long on the upper branches; stipules 8-18 mm long, the small- est at the base of the branches, the longest above, subulate-attenuate, connate 5-10 mm; leaflets 7-9, lanceolate, the tips acute, mucronate, the largest leaflets 2.5-3.5 cm long, 5 mm wide, sparsely kinky-villous above, canescent to kinky- villous below; peduncles 3 cm long at anthesis, 3-8 cm at fruiting; racemes 3-6 cm long, verticillate; bracts subulate-attenuate, 8-14 mm long, canescent; pedi- > u T. paris x Lupinus muelleri drawn to the mean nu on the scale shown, oer the. stipules and leaflet which are half scale. The lettering i a 1 Yan dorsa view; Br — bract; Ca — pom inside view; F — flower, left side L = leaflet; S = stem; St = stipule; W = wing. (See legend for Fig. 2 i. “full a. ) FIGURE 8. The 5 structures for Lupinus montanus n montanus var. austro- volcanicus are shown in the upper portion for comparison with L inae The stip- ules and leaflet are gon at half the scale. The floral traits are the same as shown for L. montanus subsp. montanus var. montanus in Fig. 2. The lower portions shows the struc- tures of L. kellermanianus drawn to the mean values on the scale shown. The stipules and leaflet are drawn to half scale. The lettering is: B — banner, dorsal view; Br — bract; C — calyx, inside view; F = flower, left side view; K = keel; L = leaflet; S — stem; St = stipule; = wing. (See legend for Fig. 2 for full explanation.) Ficure 9. Structures ua a valerioi drawn to the mean i at the das shown, except the stipules and lea which are half scale. e lettering is: B — bann , dorsal view; Br — bract; Ca — ecl inside view; F — er, left nu view leaflet; S — stem; St — stipule; W — wing. (See 3 for Fig. 2 for full pier nec ) * DUNN & HARMON—LUPINUS 361 1977] SSS — — — — —— == z === =a > =< = Le = — — == > = =a === Sa = — Lupinus muelleri Standl. [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN 362 "uS do snuDiUDUI49||9 snuidn A os CL BENT aD ů — Ts x je CAN N 2 SSS OS T — - ; A — Z = — — eZ — — * A XIS ‘WS d'2) SNIIUDIJOAOILSNO oA 'Y'G'H Snupiuou snuldnq : = s —— AR === mul —— =— „FFF P —— s —— — DUNN & HARMON—LUPINUS 363 1977] — — —S 364 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 cels 5 mm long at anthesis, 10-12 mm in fruit, hispidulous; calyces canescent, the lower lip 7.5-8.5 mm long, strigose within on the distal half, the upper lip 6.6- 6.8 mm long, the notch 1.2-1.4 mm deep, the lips connate 1.5 mm laterally, the bracteoles lanceolate, 3-4 mm long, attached well below the lateral sinus lip on the side of the calyx cup or at the base; banner with a sparse patch of kinky hairs distally on the dorsal side, densest on the crest, obcordate to suborbicular, 12 mm long, 11.5-12 mm wide, reflexed near the midpoint; wings glabrous, 12-14 mm long; keel glabrous, 44.5 mm wide in the middle, the acumen very short; ovules 6-7; pods 3.5 cm long, 7 mm wide, sparsely pilose; seeds not available. Very few collections have been made of this species known only from Volcán Agua and Zunil. The traits of the woody stems and pubescence on the banner have shown up on several neighboring peaks in plants which are otherwise typi- cal L. montanus. This suggests introgression and the material named L. mon- tanus var. austrovolcanicus is probably of hybrid derivation. UATEMALA. QUEZALTENANGO; Summit of Volcán Zunil, Steyermark 34848(F). SACA- TEPEQUEZ: Volcán Agua, 9,000 ft, Kellerman 4746( US); Kellerman 15, 1905( US). 5. Lupinus valerioi Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 545. 1937. tyre: Costa Rica, San José, Cerro de las Vueltas, Standley & Valerio 43668 (holotype F ).—Fic. 9. Plants perennial, 6-9 dm tall, woody below; stems of current season hollow, subfistulose, to 5 mm in diameter, with abundant spreading pilose hairs mm long, and with an undercoat of appressed strigose hairs; petioles of mature leaves, 6-12 cm long, pubescence as on the stems; stipules 2.5-4.5 cm long, connate to the petioles 1-2.5 cm, pilose, the free portion slender subulate-atten- uate; leaflets 7-10, slenderly oblanceolate, the largest 4-7 cm long, 7-10 mm wide, glabrous above, subappressed strigose below; foliage dense from short internodes causing the lower stipules to be imbricated on the branches, 2-3 cm, with multiple leaves from the lateral buds of the upper nodes; peduncles 8-10 cm long; racemes 6-17 cm long, those of the branches shorter, verticillate to subverticillate, the whorls 10-25 mm distant in age; bracts caducous, lance- attenuate, 15-23 mm long, 2-3 mm wide in the lower portion of the raceme, with numerous pilose hairs 2-3 mm long dorsally; pedicels 3-4 mm long at anthesis, spreading pilose, the hairs 1-2 mm long; calyces densely subappressed pilose, the hairs 1-2.5 mm long, the lower lip 8.5-10.6 mm long, the tip bi- or trifid, the teeth 0.1-0.3 mm long, the upper lip 6-8.5 mm long, bifid, the notch 3.5-5.5 mm deep, the lips connate laterally 1.8-2.4 mm, the bracteoles mm long, attached on the calyx cup below the lateral sinuses, with the lower portion fused to the calyx cup; banner glabrous, suborbicular, somewhat con- stricted below into a broad claw, 12-14 mm long, 11-12.5 mm wide, reflexed 5.5-6.5 mm, appressed 6.5-7 mm, reflexed/appressed ratio 0.76-0.83; wings 13.5-16.5 mm long, 6.5-8 mm wide, the claws 2.8-3.4 mm long; keels generally minutely ciliate above on the distal part, 3.5-4.5 mm wide in the middle, the angle 90°-95°; ovules 5; pods 3-3.5 cm long, 8.5-9.5 mm wide, densely villous, the hairs 2-3 mm long; seeds 4.5 mm long, 3 mm wide, dark brown to black. 1977] DUNN & HARMON—LUPINUS 365 The stipules and bracts show derivation from the L. montanus genome, which apparently was introduced and introgressed with a local taxon which at the present time appears to have dominated most of the characteristics, with only the vestige of traits from L. montanus. The flowering and fruiting mate- rials have been collected from September through January at elevations from 2,700-3,100 m. Costa RICA. SAN josé: Cerro Chirripó, Evans et al. 118( MICH). Cerro Frio, Jiménez 2677 (CR, F). Cerro Vueltas, Standley & Valerio 43668( F), 43974(F, US). Cord. Talamanca, Weber 6257 ( MICH ). LITERATURE CITED BEAMAN, J., C. D DE Joxc & W. P. SrovrAMiInE. 1962. Chromosome studies in the al- pine d subalpine floras of Mexico and Guatemala. Amer. J. Bot. 49: 41 Cuo, Y. O. . O. MARTIN. 1971. Resolution and 1 nambiguous identification of micro- gram m of 22 lupine alkaloids by seq: 1 use o in-layer and gas-liquid chromatography and mass spectrophotometry. Anal. Biochem. 44: Dunn, D. B. 1972. Lupinus mexicanus Cerv. ex Lag. Rhodora 74: 489—494. HorMcnREN, P. K. & W. Keuken. 1974. Index Horberiornn. Ed. 6. Regnum Veg. 92: —397. Huwmpoutpr, F. H. A. J. A. BOINPLAND & C. S. Kuntau. 1823. Nova Genera et pecies Plantarum. Vol. 2 Paris. Lacasca, M. 18 Genera Species 1 i Madric Nowacki, E. 1963. Proc. XI Int. Congr. Gen The Hapu. Vol. D) Dunn. 1964. irk Califor lupines and n suggested by alkaloidal content. Genet. Polon. 5: 47— SHARMA, G. 1967. Cuticular variation in e and Datura. Ph.D. thesis, Univ. Mis- sami-c olumbia, Columbia, Missouri. SMrru, C. 1948. Correlation-key to group 9, montanus complex. Species lupinorum. — 20 GUAYANIA DAVIDSEI AND HEBECLINIUM GENTRYI, NEW SPECIES FROM NORTHERN SOUTH AMERICA (EUPATORIEAE—ASTERACEAE )' RoaBERT M. KING? AND HAROLD ROBINSON? ABSTRACT Descriptions and discussions of relationships are provided for Guayania davidsei R. M. King & H. Robinson from the Amazonas region of Venezuela and Hebeclinium pa R. M. King & H. Robinson from the Chocó region of Colombia. Collecting efforts of two members of the staff of the Missouri Botanical Garden have provided two new species belonging to the Hebeclinium complex of the tribe Eupatorieae (King & Robinson, 1971a, 1971b). The complex is notable for the partially deciduous subimbricate involucral bracts, the simple structure of the style base, the smooth corolla lobes, the long nonannulated anther collars, and for a tendency to bear hairs on the receptacle. The latter character which was the traditional distinction of Hebeclinium is, however, not consistent throughout the group, being absent in some species of Hebeclinium sens. str. and lacking in all species of Guayania. The genera Hebeclinium and Guayania are most easily distinguished by the extremely filiform style append- ages of the former and by the strongly asymmetric carpopodium of the latter. With the present additions Hebeclinium has 19 species concentrated in the northern Andes with one species, H. macrophyllum (L.) DC., widely distrib- uted. Guayania now contains 6 species all restricted to the Cuayana Highlands region and to the surrounding lowlands of the Orinoco and Amazon. Guayania davidsei R. M. King & H. Robinson, sp. nov.—F'c. 1. Plantae herbaceae perennes erectae ca. 4 dm altae pauce ramosae. Caules succulenti anguste jatrophiformes superne abbreviati et in inflorescentia abrupte terminati glabri et in sicco irregulariter striati. Folia opposita superne congesta, petiolis 4-6 cm longis; laminae ovatae 9-12 cm longae et 6.0-7.5 cm latae pen- ninervatae base late obtusae margine serratae apice vix breviter acuminatae supra et subtus glabrae vel ad marginem sparse puberulae. Inflorescentiae laxe cymosae, ramis dense puberulis, pedicellis 0.3-1.5 mm longis. Capitula 5.0-5.5 mm longa et 3-4 mm lata; squamae involucri ca. 28 et 4—5-seriatae 1.0-4.5 mm longae ad 1 mm latae exteriores late ovatae interiores sensim oblongae vel an- guste lanceolatae margine et apice distincte scariosae et minute puberulae apice late vel anguste rotundatae extus glabrae et plerumque 3-striatae; receptacula leniter convexa glabra. Flores ca. 22 in capitulo; corollae albae tubulosae ca. 3 mm id faucis tubulosis base indistinctis, lobis ca. 0.3 mm longis et 0.25 This study was supported in part by the National Science Foundation Grant DEB77- 13457 to the senior author. ? Department of Sposa United States National Museum, Smithsonian Institution, Wash- ington, D.C. 20560 ANN. Missouni Bor. Garp. 64: 366-370. 1977. 1977 GUAYANIA AND HEBECLINIUM 367 KING & ROBINSON PLANTS OF VENEZUELA COMPOS ITAE Guayani s didi NN n e Eae dm Puerto Ay UNITED STATES elev. 1 ` Low 3 € East of highway with savanna at 2119111 " edge of la — boulder outcrops in forest shade--at least pertially--on mall hills. Heads white NATIONAL HERBARIUM Gerrit t Davidae 2845 2 November 1971 MISSOURI BOTANICAL GARDEN HERBARIUM — Ficure 1. Guayania davidsei R. M. King & H. Robinson, holotype, United States Na- tional Herbarium. Photo by Victor E. Krantz, Staff Photographer, National Museum of Natural History. mm latis, faucis superioribus et lobis extus breviter puberulis, pilis moniliformi— bus; filamenta in parte superiore ca. 0.25 mm longa; thecae ca. 0.8 mm longae; appendices antherarum oblongae ca. 0.15 mm longae et latae; grana pollinis ca 18 my in diametro; achaenia 1.5 mm longa plerumque in costis breviter setifera 368 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 setae pappi ca. 25 tenues ca. 3 mm longae superne non latiores, cellulis api- calibus argute acutis. TyPE: VENEZUELA. AMAZONAs: 20 km S of Puerto Ayacucho, 100 m, low forested hills E of highway with savanna at base, at edge of large boulder outcrops in forest in shade—at least partially—on small hills, heads white, 2 Nov. 1971, Gerrit Davidse 2845 (US, holotype; MO, isotype). Guayania davidsei is the second member of the genus with modified stem structure and abrupt bases on the inflorescences. In this species the stems are somewhat thickened with more congested leaves in the upper part, providing the superficial resemblance to a Jatropha. The related G. bulbosa (Arist.) R. M. King & H. Robinson has stems completely underground and tuberous. The latter species also differs by the glanduliferous branches of the inflorescence. The new species is apparently from low elevations, while G. bulbosa is reported from talus slopes at 1,500 m elevation on Serrania Parü. Hebeclinium gentryi R. M. King & H. Robinson, sp. nov.—F'c. 2. Plantae suffrutescentes erectae ca. 1 m altae? Caules obscure tetragoni sordide lanosi. Folia opposita, petiolis 0.5-1.0 cm longis; laminae ovatae 3.5-7.0 cm longae et 1.8-3.7 cm latae base breviter obtusae vel subrotundatae margine minute serratae vel duplo-serratae apice distincte breviter acuminatae supra virides glabrae vel glabrescentes fere ad marginem et nervis primariis minute puberulis subtus fulviores obscure glandulo-punctatae in nervis et nervulis dense sordide puberulae, nervis secondariis pinnatis valde ascendentibus paucis. In- florescentiae corymboso-paniculatae, ramis dense puberulis, ramulis ultimis 1—5 mm longis. Capitula plerumque 5 mm alta et 3.5-4.0 mm lata; squamae invo- lucri ca. 40 subimbricatae 4—5-seriatae valde inaequales 1-3 mm longae 0.3-0.6 mm latae anguste oblongae apice rotundatae margine minute dense puberulae extus plerumque trisulcatae superne in bracteis interioribus sensim dense puber- ulae; receptacula leviter convexa sparse puberula interne non scleroidea. Flores ‘a. 25; corollae albae tubulosae ca. 3 mm longae; faucis tubulosis base indistinctis glabris, lobis triangularibus ca. 0.4 mm longis et latis extus dense minute puberu- lis, pilis brevibus in apicem subclavatis; filamenta in parte superiore ca. 0.15 mm longa; thecae ca. 0.8 mm longae; appendices antherarum late oblongae ca. 0.15 mm longae et latae; grana pollinis ca. 20 mu in diametro; achaenia 1.5 mm longa sparse minute glandulifera superne pauce sed non breviter setifera; setae pappi ca. 45 plerumque ca. 2.5 mm longae ad apicem vix vel non latiores, cel- lulis apicalibus obtusis. Tyre: CoLoMnBia. cuocó: Alto de Buey, 1,200-1,800 m, tropical wet forest, weak-stemmed shrub, flowers white, 8 Jan. 1973. Al Gentry & Enrique Forero 7290 (US, holotype; MO, isotype). Hebeclinium gentryi is most closely related to H. reedii R. M. King & H. Robinson (King & Robinson, 1972) of adjacent Darién Province of Panama. The two species share the lanate stems, the pubescent outer surfaces of the involucral bracts, less hemispherical receptacles than usual in the genus, and KING & ROBINSON—-GUAYANIA AND HEBECLINIUM 369 PLANTS OF DEPARTMENT or Choc "Nm UNITED STATES 211 Alto de Buey, alt. 1200-1800 meters, 9193 tropical wet forest Al Gentry & Kamine Forero no. 5-55 NATIONAL HERBARIUM Dae 8 Jan 197) — MISSOUM! BOTANICAL GARDEN HERBARIUM {mo} Ficure 2. Hebeclinium gentryi R. M. King & H. Robinson, holotype, United States National Herbarium. leaves with at least partially doubly serrate margins. The new species does have a different appearance from H. reedii by the smaller leaves and the more neatly oblong subimbricate involucral bracts, but the more significant differ- ences are the glabrous to glabrescent upper leaf surfaces, the less pubescent 370 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 surface and larger area of pith on the receptacles, the scarcely enlarged tips of the pappus setae, and the glanduliferous achenes. In H. reedii the upper surfaces of the leaves are coarsely pilose, the receptacles are shortly but densely puberu- lous on the ridges, the pith of the receptacles is very reduced, the pappus setae have prominently enlarged tips, and the achenes are glabrous. LITERATURE CITED KiNG, M. & H. RoBINSON. la. Studies in the Eupatorieae (Asteraceae). XXXVII. T genus sap ae "Phytologia 21: -301 ——— & ———. 1971b. ies in the Furatiniene (Asteraceae). XXXIX. A new genus, Guayania. adus 21: 302—303. 1972. Studies in the Eupatorieae (Asteraceae). LXXIV. New species of Critonia, a Lapis and Hebeclinium. Phytologia 23: 405—408. NOTES A NEW SPECIES OF BAUHINIA (LEGUMINOSAE) FROM PERU Continued revisionary studies of the neotropical species of Bauhinia have resulted in the discovery of a new species of the genus endemic to Peru. Bauhinia hirsutissima Wunderlin, sp. nov. Frutex scandens cirrhosus; rami juvenales fuscoporphyro-hirsuti. Folia anguste ovata ad oblonga, ca. 1⁄4 vel rare 74 longitudine bilobata, 5-14 cm longa, 4-10 em lata, apice acumi- nata ad obtusa, basi profunde cordata, margine revoluta, chartacea ad subcoriacea, supra glabra, infra fuscoporphyro-hirsuta, 9- ad 11-nervata; petioli 3-5(—7) em longi; stipulae reni- nflores i fuscoporphyro-hirsutae; rhachis 12-40 cm longa; gemmae ovoideae, excrescentibus apicalibus gemmarum ovato-lanceolatis, 3-5 mm longis, ugs atis; bracteae et bracteolae anguste lan- ceolatae, 5-7 mm longae; pedicelli graciles, 5-10 mm longi; hypanthium oue ca. 1 mm longum calyx 5 vel leviter 15 5 a 15-nervatus; petala 5, subRequalia, alba vel subrosea, 10-12 mm longa, lamina late elliptica, intra glabra, extra dense ess pilosa, ungue lamina 3 vel subaequalia; stamina fertilia 10, tubo calycis longitudine E libra, 5 glabra, 5 versus apicem pilosa, antheris oblongis, ca. 1 mm longis; gynoe- cium staminibus longitudine plus minusve aequale, stylo brevi, crasso, arcuato 1 ovario dense hirsuto, gynophoro minuto, „ obliquo. Legumen dehiscens, oblongum ad anguste obovatum, 5 ca. 7. | longum, ca. 2.5 em latum, brunneum glabratum; semina suborbiculata, 11-12 diam., pa ds hebetata puncticulata, obire striata, brunnea, cicatricibus funiculi ramorum longitudine subaequalibus, ca. 1 mm longa. Tendriled woody vine; young branches reddish brown hirsute, glabrescent in age, older stems not seen; intrastipular tendrils single or paired, woody, cir- cinate. Leaves narrowly ovate to oblong, bilobate ca. 1⁄4 or rarely 24 their length, 5-14 cm long, 4-10 cm wide, the apex of lobes acuminate to obtuse, the base deeply cordate, the margin revolute, chartaceous to subcoriaceous, glabrous above, reddish brown hirsute below, the lower surface frequently purplish- tinged, 9-11-nerved; petioles 3-5(-7) cm long, reddish brown hirsute; stipules reniform, 5-10 mm long, 2-5 mm wide; intrastipular excrescences other than tendrils minute. Inflorescences racemose, terminal or subterminal and axillary, elongate, slender, lax, reddish brown hirsute throughout; rachis 12-40 cm long, the lower flowers soon deciduous, the inflorescence then frequently with 10-30 flowers on a long-pedunculoid rachis; buds ovoid, 6-8 mm long, the free tips ovate-lanceolate, 3-5 mm long, incurved; bracts narrowly lanceolate, 5-7 mm long; bracteoles similar to the bracts, but smaller, attached near or above the middle of the pedicel; pedicels slender, 5-10 mm long; hypanthium cyathiform, ca. 1 mm long; calyx campanulate or slightly bilabioid at anthesis, 15-nerved, each trio of nerves ending at one of 5 ovate-lanceolate appendages at the rim of the calyx tube, the median nerve extending the length of appendage, the lateral 2 ending at the base of the appendage or inconspicuously extending up to % its length; petals 5, subequal, white or faintly tinged with pink, 10-12 mm long, the blade broadly elliptic, 5-7 mm long, 3-4 mm wide, glabrous internally, densely appressed brown-pilose externally, the claw longer than to nearly equal- 379 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ling the length of the blade, brown pilose; fertile stamens 10, + equalling the calyx tube, free to the base, alternate ones slightly shorter, the longer 5 gla- brous, shorter 5 brown pilose towards the tip, the filaments arcuate, the anthers oblong, ca. 1 mm long, white-pilose; gynoecium + equalling the stamens, the style short, thick, arcuate, glabrous, the ovary densely brown-hirsute, the gyno- phore not evident, the stigma oblique, slightly differentiated from the style. Fruit a dehiscent legume, oblong to narrowly obovate, apiculate with a per- sistent style, ca. 7.5 cm long, ca. 2.5 em wide, dark brown, glabrate, gynophore not seen; seeds suborbicular, ca. 12 mm long, ca. 11 mm wide, the surface dull, puncticulate, obscurely striate, dark brown, funicular-branch scars subequal, ca. 1 mm long. Chromosome number unknown. TYPE: Peru. Loreto: Fortaleza, near Yurimaguas, ca. 140 m, forest, Dec. 1932, G. Klug 2800 (US, holotype; F, MO, NY, isotypes). Specimens examined: PERU. LoRETO: Quebrada Shanuce above Tu 71 v (MO, duplicates to be distributed). Lower Río Huallaga, Killip & Smith 27601 (F, US), 28302 (F, NY, US). Fortaleza, Yurimaguas, Ll. Williams 4216 (F, US), prt (F, i Distribution: Known only from near Yurimaguas on the Río Huallaga, Lo- reto, Peru. It occurs in forests at elevations of about 140 meters. Flowering material has been collected from July through December with nearly mature fruiting material collected in December. The newly described species is most closely related to the widespread and highly variable B. glabra Jacq., but is distinguished by the following combina- tion of characters: young branches, inflorescences, and lower leaf surfaces con- spicuously deep reddish-brown hirsute; inflorescences elongate, slender, lax, with flowers distantly arranged; flower buds ovate, with incurved lanceolate apical excrescences; petal blades 5-7(-8) mm long, without purple spots. In contrast, the vestiture of B. glabra, when approaching that of B. hirsutissima, is pilose and has a coppery sheen. One local race of B. glabra whose pubescence is very similar to that of B. hirsutissima is restricted to Panama and differs in all other respects. The flowers of B. glabra have petal blades 10-20 mm long, one of which is usually conspicuously marked with purple spots, although some- times obscure in local races. In B. glabra the flower buds are lanceolate with setiform or rarely lanceolate apical excrescences and the inflorescences, if elon- gate, are strict and with the flowers more closely arranged. Bauhinia reflexa Schery, a Panamanian and Colombian species, has the vesti- ture of its leaves, young branches, and inflorescence rachis in addition to the purple-tinged undersides of its leaves like B. hirsutissima, but differs in all other respects. Finally, B. hirsutissima also superficially resembles B. killipiana Standley and to a lesser degree B. vulpina Rusby and B. porphyrotricha Harms, but dif- fers slightly in nearly all characters. Specimens determined to be B. hirsutis- sima have frequently been identified by other workers as B. porphyrotricha. Examination of type material of B. killipiana, B. vulpina, and B. porphyrotricha revea] that these species are best placed in synonymy with B. glabra sensu lato. 1977] NOTES 973 I gra itefully acknowledge John D. Dwyer, St. Louis University and Missouri 1 Garden for critically reading the manuscript, especially the Latin description, and the cura- tors of the herbaria who pee specimens for this study —Richard P. Wunderlin, Department of Biology, University of South Florida, Tampa, Florida 33620. ALNUS MARITIMA MUHL. EX NUTT., NOT ALNUS METOPORINA FURLOW The new name, Alnus metoporina Furlow, proposed in a recent issue of this journal ( Furlow, 1976) to replace the long-recognized Alnus maritima Muhl. ex Nutt., is unnecessary according to Article 55 of the International Code of Botani- cal Nomenclature (Stafleu et al., 1972) which states: When a species is transferred to another genus or placed under another generic name dd the same genus without change of rank, the specific epithet, if legitimate, must be retained "s has not been retained, must be reinstated unless one of the following obstacles exists: The resulting binary name is a later homonym 64) or a tautonym (Art. 23). T An earlier legitimate specific epithet is available (but see Arts. 13f, 58, 59, 72). =< > E — The genus Betula-Alnus was validly published by Humphry Marshall in his Arbustrum Americanum (1785), and three species, including B. maritima, placed in it. Marshall's description of B. maritima is sketchy, mentioning only the height of the plant, the Jong and narrow” leaves, and the very distinctive August anthesis. Marshall did not cite a type nor did he keep an herbarium, and so we do not know on what material the species was based. However, we have no doubt that the plant was the same species that was later described by Henry Muhlenberg in an unpublished manuscript, and subsequently validly published by Thomas Nuttall (1842) in the first volume of his Sylva. Alnus maritima was described as a new species in the genus Alnus, based not upon Marshall's name [as assumed by some authors, see Little (1953)] but upon Muhlenberg’s manuscript name. We know that Muhlenberg was well acquainted with Marshall’s work for in a letter to William Bartram dated 10 December 1792 (Darlington, 1849), he wrote: “Marshall has given me some satisfaction, but his Arbustrum wants some emendations. Any observations that way where you think he is wrong, or where another name might have been given, would be so pleasing to me.” It is likely that Muhlenberg may have recognized that his species was Betula- Alnus maritima of Marshall, and that he intended to make the transfer to Alnus, and to typify the species on the Bartram collection he had. However, there is nothing in the manuscript preserved at the Philadelphia Academy of Sciences Library to suggest this, and we may only speculate on Muhlenberg’s intentions. Nuttall apparently put even less faith in Marshall’s work, for he never men- tioned it directly in the first volume of his Sylva. Marshall is cited there only once, and then in synonymy under Nuttall’s Carya microcarpa. Since a Muhlen- 374 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 berg synonym is also cited, it is possible that the Marshall reference was copied by Nuttall from Muhlenberg. Furlow (1976) is correct in his contention that to transfer Marshall's earlier epithet from Betula-Alnus to Alnus would create a later homonym ( prohibited by Articles 55[1] and 64). His creation of a new name for B. maritima to avoid the creation of a homonym, however, is not correct. Article 55(2) requires that if an earlier legitimate specific epithet is available for the species, it must be adopted. Alnus maritima as proposed by Nuttall (1842) is legitimate, is avail- able, and antedates Furlow's A. metoporina. Alnus maritima Muhl. ex Nutt. must be retained for this plant. We propose to lectotypify Alnus maritima on the Pickering specimen (PH) on which Nuttall’s (1842) description and Plate X (bis) were based, and not on the Bartram specimen (#477 in Herb. Muhlenberg, PH) which was also seen and indirectly mentioned by Nuttall. Furlow has supplied a neotype for Marshall's Betula-Alnus maritima. We would like to thank E staff at the Philadelphia Academy of Sciences for their courtesies, and Dr. Dan H. Nicolson of the National Museum of Natural PEN Smithsonian Institution, for his advice on the provisions of the Code discussed herein LITERATURE CITED DanLiNGTON, W. 1849. Memorials of John Bartram and Humphry Marshall. Lindsay & Blakiston, Philadelphia. FunLow, J. : Nomenclatural changes in Alnus (Betulaceae). Ann. Missouri Bot. Gard. 63: 380-381. LrrrLE, E. L. 1953. Check list 5 native and naturalized trees of the United States (includ- ing Alaska). Agric. Handb. 41: 1-472. MansHALL, H. 1785. b LE The American Grove. J. Crukshank, Phila- phia. NurTALL, T. 1842. The 1955 American Sylva. Vol. 1. Smith & Wistar, Philadelphia. SrAFLEU, F. A. ET AL. (editors). 1972. International code of botanical nomenclature. Reg- num Veg. 82: 1-426. —Virginia M. Stibolt, C. Rose Broome and James L. Reveal, Department of Botany, University of Maryland, College Park, Maryland 20742. NEW TAXA AND COMBINATIONS IN THE GENUS LASIACIS (GRAMINEAE) The following new taxa and new combinations are published in advance of my systematic treatment of the genus Lasiacis in order to make them available for use in other publications. A new species, Lasiacis nigra Davidse, has also been published (Davidse, 1974). Full explanations for my treatment will be given in the forthcoming publication. Lasiacis divaricata (L.) Hitchc. var. austroamericana Davidse, var. nov. Ab L. divaricata var. divaricata spiculis globosis et ramis paniculae ascen- dentibus differt. 1977] NOTES 375 Tyre: BRAZIL, MINAS GERAIS: Near Santa Barbara do Caparaco, streamside, suffrutescent, 3 m high, climbing and rooting at lower joints, “canaviera,” 21 Nov. 1929, Mexia 4007 (NY, holotype; F, GH, UC, US, isotypes). Lasiacis divaricata (L.) Hitchc. var. leptostachya (Hitche.) Davidse, comb. et stat. nov. Lasiacis leptostachya n Contr. U.S. es Herb. 22: 19. 1920. Type: NICARAGUA. Jinotepe, jungle, 500 m, stout central canes, branches more or less whorled and ed floral branches odia n flexuous, pce all small, 7 Nov. 1911, Hitchcock 8718 (US, holotype). Lasiacis grisebachii (Nash) Hitche. var. lindelieana Davidse, var. nov. Ab L. grisebachii var. grisebachii laminis latioribus (1.5-2.0 cm) differt. Type: CUBA. HABANA: Lomas de Camoa, in siloa locis umbrasis satis fre- quens, 27 Nov. 1921, Ekman 13530 (US-1295003, holotype; F, NY, US-1502317, isotypes ). Lasiacis oaxacensis ( Griseb.) Hitchc. var. maxonii (Swallen) Davidse, comb. et stat. nov. Lasiacis maxonii idcm Ann. Missouri Bot. Gard. 30: 231. 1943. TYPE: PANAMA. CHIRIQUI: Vicinity of El Boquete, in thickets Mode i^ in ail, 1,000-1,300 m, 2-8 Mar. 1911, Maxon 4999 cds holotype; US, isotvpe). Lasiacis rugelii (Griseb.) Hitchc. var. pohlii Davidse, var. nov. Ab L. rugelii var. rugelii spiculis globosis minoribus, inflorescentiis minori- bus, ramis paniculae ascendentibus vel patentibus, pseudopetiolis amplis differt. Type: Costa Rica. carraco: 1 km NE of Pejibaye, along Río Pejibaye, growing at base of tree, ca. 700 m, 2 Nov. 1968, Pohl & Davidse 11478 (ISC, holotype; CR, EAP, K, MO, US, isotypes). Lasiacis ruscifolia ( H.B.K.) Hitchc. var. velutina (Swallen) Davidse, comb. et stat. nov. Lasiacis velutina Swallen, Ceiba 4: 288. 1955. Type: HONDURAS. MORAZÁN: Vicinity of El Zamorano, road to San Antonio, 17 Oct. 1951, Swallen 10834 (US, holotype). Lasiacis sorghoidea (Desv.) Hitchc. & Chase var. patentiflora (Hitchc. & Chase) Davidse, comb. et stat. nov. Lasiacis patentiflora Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 338. 1917. Type: Tonaco: > f island, edge of woods on mountainside, 20 Dec. 1912, Hitchcock 10268 (US- 865566, holotype; US-975660, isotype) LITERATURE CITED Davipse, G. 1974. A new species of Lasiacis (Gramineae). Phytologia 29: 152-153. —Gerrit Davidse, Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. 376 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 SOLANUM ARMENTALIS: A NEW SPECIES FROM COSTA RICA Solanum armentalis J. L. Gentry & D'Arcy, sp. nov.—Fic. I. Frutex 1 m altus inermis omnino confertim stellato-pubescens, trichomatibus fulvis, sessili- radio medio quam radiis lateralibus multo longiore; foliis singularibus vel in geminis — PS — - — = — n O — — =) > — — n — = [e] 7 [e] = > '" " 1 > O = = = = — = w = = = = = ^ Q — — = = = vel ovato-ellipticis, petiolo 5-7 mm longo; foliis minoribus ovatis vel late ellipticis; i 8 lateralibus foliis opposita, pedunculo supra 5 cm bifurcato; floribus parvis, calyce 2-2. mm longo prope basim fisso, lobis ovatis, 1.5-2.2 mm longis, apice acutis, staminibus aequalibus, filamentis glabris, basim connatis, antheris = mm longis, poris largis terminalibus aperientibus, ovario glabro, stylo glabro; acino ignote Shrub 1 m tall T densely stellate-pubescent throughout, the hairs sessile, yellowish brown with the central ray greatly exceeding the lateral rays. Leaves solitary or in pairs, unequal in size, similar or different in shape, densely pubescent beneath, sparsely pubescent above except along the midvein; larger leaves ovate to ovate-elliptic, the blade 13-14 cm long, 6-7 cm wide, the apex long acuminate, the base rounded, the petiole 5-7 mm long; smaller leaves ovate to broadly elliptic, the blade 1.8-4 cm long, 1.2-2.8 cm wide, the petiole 2-3 mm long. Inflorescence both lateral and opposite the leaves, several flowered; pedun- cle unbranched for 5 cm, bifurcate; pedicels slender, 10-12 mm long. Flowers small; calyx 2-2.5 mm long, parted to near the base, the lobes ovate, 1.5-2 mm long, the apex acute; corolla white, the limb 10-11 mm wide, parted to near the base, the lobes 4-4.5 mm long with sessile stellate pubescence externally; stamens equal, the filaments glabrous, ca. 0.7 mm long, basally connate, the anthers 2. mm long, the pores large; ovary glabrous, the style glabrous, exceeding the sta- mens. Fruit unknown. TYPE: Costa RICA. PROV. PUNTARENAS: Open forest 1 mi due south of San Vito de Java, ca. 3500 ft., 18 Aug. 1967, Peter H. Raven 21887 (MO-2304198, holotype; F, isotype). Additional collection examined: Cosra Rica. prov. SAN JOsÉ: Vicinity of El General, edge of forest, 700 m, Jan. 1939, 1 F. Skutch 3919 (MO) Solanum armentalis is recognized from other species of the genus in the Central American flora by the softly tomentose leaf undersides, long, slender peduncles and slender pedicels, and small white flowers which have calyces divided nearly to the base. The indumentum is of dense, sessile stellate hairs, with the midpoints many times longer than the several short lateral arms. This species is a member of subgenus Brevantherum. However, it is not easily placed in a section within the subgenus. At first glance the plant is sug- gestive of Solanum extensum Bitt. and S. cordovense Sessé & Mog., but the calyx is different in shape and not accrescent to judge from the one immature fruit on the Skutch specimen. Also, the indumentum and inflorescence are different. Solanum gemellum Sendt. and S. megalochiton Mart. of eastern Brasil have caly- ces similar in shape, but they are conspicuously accrescent in fruit. While the above similarities are suggested, they do not imply a close relationship to the species described here. 1977] NOTES 377 Ficu Solanum armentalis J. L. Gentry & D'Arcy. B. Stem dete stellate hairs (3).—C. Flower (214). —A. Flowering branch (»x!5).— 72 [After Raten 21887 (MO). 378 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 It has been collected twice in Costa Rica, at 700 and 1,165 m elevation on the Pacific side of the isthmus. Perhaps further collections will provide an insight into its ecological requirements. The locality of San Vito suggests that this spe- cies may be found in the nearby Chiriquí Mountains of Panama. Because no single character delimits this well-marked species, the specific epi- thet does not relate to the collector, collection locality, or morphological features. —Johnnie L. Gentry, Jr., Bebb Herbarium, Department of Botany-Microbiology, University of Oklahoma, Norman, Oklahoma 73019 and William G. D'Arcy, Mis- souri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. HERBERTIA (IRIDACEAE) REINSTATED AS A VALID GENERIC NAME The genus Herbertia was described in 1827 by Robert Sweet for a New World genus of Iridaceae. As currently circumscribed (Goldblatt, 1975), the genus is a small one of approximately six species centered in temperate South America from Uruguay to Chile with a subspecies of a South American taxon occurring in the southern United States. The existence of the similar name Herbertus Gray (also used in the form Herberta) published in 1821, prompted several authors including myself to regard Herbertia as a later homonym and therefore to reject it. Following Kuntze (1898) who first suggested that Herbertia be considered a homonym, both Foster (1945) and Ravenna (1968) among others, accepted Alophia Herb. ( dat- ing from 1840) as the valid name for the genus. Subsequently I discovered ( Goldblatt, 1975) that the type species of Alophia had been misinterpreted and was in fact a species of what was then known as Eustylis. I therefore proposed another available synonym of Herbertia, namely Trifurcia Herb., for the genus and provided new combinations in Trifurcia for both the United States subspe- cies and for the South American representatives of Herbertia. Recently it has been suggested both in print (Florschutz & Grolle 1975) and to me personally that Herbertia should not have been rejected and that I should carefully consider Article 75 of the Botanical Code of Nomenclature. This article deals with names of similar but not identical spelling, and recommends rejection only in cases of likely confusion. The article recommends the rejection of exam- ples such as Columella and Columellia and Eschweilera and Eschweileria as being too similar and thus likely to cause confusion. Other examples are Pel- tophorum and Peltophorus, Iria and Iris, neither of which are to be considered homonyms and therefore both forms of these words are available for usage for different genera. Herbertia seems to fall into the latter category, being sufficiently different in orthography to avoid any possibility of confusion. Florshutz & Grolle (1975) support this view, as do several colleagues with whom I have discussed the 1977] NOTES 379 question. Therefore, I propose reinstatement of Herbertia and the synonymizing of Trifurcia. This treatment involves some new combinations as follows: l. Herbertia lahue (Molina) Goldbl., p nov. Basionym: Ferraria lahue, Molina, Sagg. Stor. Nat. Chile, ed. 2 1810 The subspecies of this taxon are to be cited as follows: la. H. lahue subsp. amoena (Griseb.) Goldbl., comb. nov. Basionym: Her- e amoena Griseb., Abh. Königl. Ges. Wiss. Gottingen 24: 325. 1 ahue subsp. caerulea (Herb.) Goldbl., comb. nov. Basionym: Trifur- cia — Herb., Bot. Mag. 1840: tab. 3779. 0 Herbertia tipridioides (Hick.) Goldbl, comb. nov. Basionym: Alophia tigridioides Hick., Darwiniana 1: 116. 1924. bo The four remaining species were either originally placed in Herbertia, or have in the past been transferred to the genus. These are: H. pulchella Sweet, H. amatorum C. H. Wright, H. hauthallii (O. Kuntze) K. Schum, H. brasiliensis Baker. LITERATURE CITED FLonscHuurz, P. A. & R. GnoLLE. 1975. He drip Gray 1821, Herbertia Sweet 1827 and Herberta Gray mut. Lindb. 1875. ]. Bryol. 8: 479-481. Foster, R. C. 1945. Studies in the 1 III. Contr. Gray Herb. 155: 3-55. GOLDBLATT, P. 1975. Revision of the bulbous Iridaceae of North America. Brittonia 27: 5. Kuntze, C. E. O. 1898. Revisio Generum Plantarum. Vol. 3. Leipzig. “pais? P. 1968. Notas sobre Iridaceae. HI. Bonplandia 2: 273-291. —Peter Goldblatt, B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. The previous issue of the ANNALS OF THE Missouri BOTANICAL GARDEN, Vol. 64, No. 1, pp. 1-143, was published on 26 July 1977. zy E Seki of Moraea (Iridaceae) in T T Africa Peter Goldbi I A New Classification of Ficus William unies |: cl s ss Studies in sess S 25: New Species and Combinations in E^ : American Bignoniaceae Alwyn H. Gent New Records of Apocynaceae for Panama and the Chocó Alwyn H. co 1 | : ‘New’ Taxa and Fonsi atibus in Eragrostis (Poaceae) John T. 1 5 Realignment o of the e Species Placed in nes (Convolvulacea š J : Aust u Aud : a The Lanes montanus Don of Mexico and Central America D B. Dun ve WU S Harmon m ĩðù ae | x .. Guyania davidsei and Hebeclinium gentryi, New Species from Northe ire pores merica ee eee Robert M. Kin, ng e ob : NOTES A New Species of Bauhinia (Leguminosae) from Peru Richa Wunderlin 9 mariti ma Muhl ex Nutt., not Alnus metoporina Pane Virginia Stibolt, C. Rose Broome d» James L. Reveal New Toa and Combinations i in ne Genus. Lasiacis (rraipen) Ge : Solanum mental | A A, Now Gent me SSOURI BOTANICAL GARDEN VOLUME 64 1977 ee e NUMBER 3 B. L usss > usa SEIWA-EN JAPANESE GARDEN, MISSOURI BOTANICAL GARDEN CONTENTS Systematics of Oenothera Sect. Kneiffia (Onagraceae) Gerald B. Straley .. 381 The South American Species of Oenothera Sect. Oenothera eens Ren nneria; Onagraceae) Werner Dietrich — Wood Anatomy of Onagraceae: Additional Species and Concepts Sherwin Carlquist Sm P t 5 ee A New a of Lopezia Loreen 2 from Sinaloa, Mexico Peter . a I. iue tru siente 8 Reinterpr m of the Type of Godetia bottae FX * Peter H. Raven & Dennis R. Parnell . „ Reproductive Structures and Evolution in Ludwigia e Ker L An droecium, Placentation, Merism Richa rd H. Eyde . SP ME W. T Dies Editor—Flara F Panama Missouri Deni pui xar len A Botanical Garden Tom Published four times a year by the St Leuk Missouri 63110. For subscription information contact. the B usin bes nec of the Annals, P.O. Box 368 1041 New Hampshir re, Lawrence, Kansas 660. cem ie price is $40 pe r volume 5 esp Senge and Meer ° $45 all I other countries. Four es volume Second clas ANNALS OF THE MISSOURI BOTANICAL GARDEN VOLUME 64 1977 NUMBER 3 SYSTEMATICS OF OENOTHERA SECT. KNEIFFIA (ONAGRACEAE)! GERALD B. SrnALEY? ABSTRACT Based on cytology, morphology and field studies, a reevaluation of Oenothera sect. k. the eastern North American day- qi odd or sundrops, is presented, with five species recognized. Subsect. Peniophyllum contains a single annual species, O. lindfolia, of the southeastern United States, which is a self-compatible diploid (n — Subsect. Eukneiffia contains one annual self- -compatible diploid (n — 7), O. d largely of the southern Mississippi Valley, an ree perennials, O. perennis, O. fruticosa, and O. pilosella. Oenothera perennis is a self-compatible Syed (n= = 7), complex heterozygote, forming a ring of 14 chromosomes at meiotic metaphase I and having about 50% pollen s terility. It is dis- tributed from Newfoundland to Manitoba and south to North Ca and Missouri. The two other perennials, each with two subspecies, are self-incompatible polyploids that seem to lack chromosomal translocations but form numerous rings of chromosomes at meiotic metaphase I owing to their autopolyploidy. Oenothera pilosella subsp. pilosella, a taxon largely of the 5 United States, is known only as an octoploid (n = pu M wk pilosella ii sessilis, also octoploid, is treated as a second subspecies. It i at present and i found in remnant prairies of the lower Mississippi River Valley. 1 “differs 880 Bbw. Pow *I am especially grateful to Peter H. Raven for originally suggesting this project and for his untiring encouragement throughout the course of this study. This study has been sup- orted in part by grants from the U.S. National Science Foundation to him. I am also grateful to the staff of te Missouri Botanical Garden for their assistance during the summer of ; where this project was completed. I would like to thank Ro M. Llo his encourage- ment and assistance, especially during the initial stages of this study, as a part of my M.S. degree research at Ohio o University. I would also like to thank Charles W. Hagen, Adolph Hecht, Brij M. Kapoor, and Lytton J. Musselman for helpful suggestions, contributions of unpublished information, and he assistance; Peter Hoch for taking the SEM photographs; Julie Wilson for assistance with the maps and lates: and Lesley Bohm for the drawings ( Figs. 81, 82). I was assisted in io field by John Ayers and Edward Minnick and I would like to thank them for their assistan Lastly, I would like to Dat the staffs of the following herbaria for the use of their herbaria during the course of this study and for the loan of material for study: BH, BHO, BM, CAN, CM, CU, DAO, DUKE, F, FLAS, FSU, GA, GH, IA, ILL, ILLS, ISC, KE, LINN, MASS, MICH, MO, MSC, NA, NCU, NEBC, NFLD, NO, NY, OKLA, OKL, OS, P, PAC, PH, POM, RSA, SIU, SMU, TENN, TEX, TRT, UARK, UBC, US, USF, VDB, VT, VPI, WIS, WVA, YU, and Louisiana Tech University, ipio College, Old Dominion Univer- sity, St. Mary's University, and Western Kentucky University. 5 of Botany, University of British Columbia, Vancouver, British Columbia V6T 1W5, Canada. ANN. Missouni Bor. Garp. 64: 381—424. 1977. ~ 382 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 — in being nonrhizomatous; it has narrower, ascending leaves, more ellipsoid, nearly sessile cap- 14) and hexaploid (n — 21), and a more northern and higher elevation subspecies, subsp. glauca (O. tetragona Roth), which as far as known is uae tetraploid. The Onagraceous genus Oenothera has been the subject of classical studies of reciprocal translocations and their effects on the evolution of a group of plants. Much of the early cytogenetic and cytotaxonomic work was done on the North American species of sect. Oenothera (Cleland, 1972). Other portions of the genus have received less attention, but studies of Raimannia (Hecht, 0), of several subgenera by Hagen (1950) and Gregory & Klein (1960), and of the South American species of sect. Oenothera by Dietrich (1977) are among the most important works. Subgenera are treated in this paper as sections consis- tent with the classification employed elsewhere in the family (Lewis & Lewis, 1955; Raven, 1964; Raven & Gregory, 1972b). Oenothera sect. Kneiffia has received little biosystematic attention. Taxo- nomic treatments of Kneiffia based largely on morphological characters have resulted in a confusing array of specific, subspecific, varietal, and form names. Even the most recent, rather thorough revision based upon morphological fea- tures by Munz (1965) reflects the difficulties in circumscribing species and particularly in delimiting infraspecific taxa. A recent biometrical and biochem- ical study (DeTurck, 1969) has also failed to clarify the taxonomic problems in this section. Records of chromosome numbers and meiotic pairing in species of sect. Kneiffia are few and scattered in the literature, and many of these, partic- ularly the older records, are from plants grown in botanical gardens for which the original source of material is not known, and for which no voucher specimens were made. There are no previous reports of complex heterozygosity in sect. Kneiffia, although polyploidy has been reported a number of times, especially by Gregory & Klein (1960) and by Laws (1963). Plants in this section are characterized by bright yellow flowers that open after sunrise, relatively short floral tubes, and 4-angled to 4-winged fruits borne on a short stipe. They are distributed from Newfoundland to northern Florida and westward to southeastern Manitoba and eastern Texas. Natural populations usually have easily discernable limits, particularly in the perennial species, and are commonly composed of a few to rarely more than several dozen plants. The two annual species tend to be found in scattered populations with less distinct boundaries. MATERIALS AND METHODS This study was begun in 1973, initially in an effort to evaluate the cytology of the section as a basis for understanding the morphological discontinuities, particularly among the perennial species. Extensive field studies were made from 1973 to 1976 throughout much of the distribution of the section. Vouchers are deposited at MO. Flower buds were fixed in the field in 1:3 glacial acetic acid : ethanol and 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 383 later transferred to 95% ethanol and refrigerated. The buds were hydrolyzed in 1:1 concentrated hydrochloric acid : 95% ethanol at room temperature until they were translucent ( 10-20 minutes depending on bud size) to remove starch grains from the pollen mother cells (Lewis & Lewis, 1955). After hydrolysis, the anthers were removed, stained with aceto-carmine and macerated with iron needles. A drop of Hoyer’s medium (Beeks, 1955) was added to make the slide permanent. The preparation was squashed and later the coverslip was ringed with Diaphane. Observations of meiotic divisions of microsporocytes were made during diakinesis or metaphase I using a positive phase-contrast microscope. Pollination systems and floral biology were studied in several natural popu- lations, and seeds or plants were collected and grown in the greenhouse or experimental garden for compatibility studies and further observation of charac- ters. Field collections have been supplemented by examination of over 11,000 herbarium specimens, including large enough samples of all taxa to be of great value in comparative morphology. Where available, herbarium voucher speci- mens and permanent slides of cytological material from previously reported chromosome studies of sect. Kneiffia were borrowed for further study. Based on a reevaluation of morphological characters and the results of cyto- logical and breeding studies, a revised classification of the taxa is presented here. HISTORICAL AND TAXONOMIC CONSIDERATIONS Due to their widespread distribution and abundance in eastern North Amer- ica, the species of sect. Kneiffia have been the subject of study for more than 200 years. Linnaeus described the first species, Oenothera perennis, in 1753. The classification of this group subsequently has undergone many reorganiza- tions owing to the difficulties in assigning populations to clear-cut morphological species. The greatest taxonomic problems are in the two perennial, polyploid species treated by Munz (1965) and in most recent floras (e.g., Gleason, 1963; Fernald, 1950; Radford et al, 1968) as O. fruticosa L. and O. tetragona Roth, including O. glauca Michx. [= O. fruticosa L. subsp. fruticosa and O. fruticosa L. subsp. glauca ( Michx.) Straley, respectively, in this treatment]. ; In early studies of the section (e.g., Michaux, 1803; Pursh, 1814) O. glauca, with broader leaves, glandular hairs, and oblong capsules, was considered dis- tinct from O. fruticosa, with narrower leaves, nonglandular hairs, and distinctly clavate capsules. At this time, there was a paucity of herbarium material avail- able for study and few, if any, intermediate populations had been sampled. Later, as more material became available, Spach (1835), Torrey & Gray (1838- 1840), Small (1896), and Léveillé (1902), in accord with the trends of the times, assigned taxonomic status to more of the intermediate populations, giving them specific, varietal, or form rank. Watson (1873), however, took a more conserva- tive view, recognizing only O. fruticosa and O. glauca. Pennell (1919) in the first thorough study of the group, recognized nine species with three varieties within this species complex. He was clearly not completely satisfied with his taxonomic treatment, however, as shown by his statement, “I present the results of this study with hesitation. Species lines have not always been found clear, 384 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 and in any genus so near to Oenothera one may expect the same tendency to split into incipient species" (Pennell, 1919: 363). Munz (1937) simplified the classification of this group by recognizing only the two species in the complex, but he retained 12 infraspecific entities; in 1965, Munz still recognized only two species, with 10 infraspecific entities, and stated that the two were in need of cytological and experimental work. In regional floras where Munz's treatments are followed closely, comments are often made concerning the variability of the two species (e.g., Fernald, 1950; Gleason, 1963). Radford et al. (1968: 752) say of O. tetragona, "This species complex and number 9 [O. fruticosa] are poorly understood, both are in need of biosystematic study." W. J. Hooker (1837, sub pl. 3545) was far ahead of his time when he astutely observed that “O. fruticosa is a species, widely extended throughout North America from Canada to Carolina; but so variable in its foliage and hairiness, as to suggest the idea of there being the several species above enumerated.” In this species he included 10 previously recognized taxa, even the broad-leaved O. glauca. At this early date Hooker recognized the variability within this species complex and its essential cohesiveness. Thus two philosophies have emerged in regard to the systematics of this complex. The one, splitting and assigning formal taxonomic status to many pop- ulations, contrasts with the other, lumping populations into taxa with broad species lines. Difficulties in splitting the section into a large number of taxa are caused by the infraspecific variability. Individual plants within one population can be assigned to different taxa, and there are countless morphological interme- diates forming a continuum from one taxon to another, making distinct narrow specific lines impractical. Consequently, the treatment here of O. fruticosa with two relatively distinct subspecies, both morphologically and geographically, seems the only practical and taxonomically sound means of reflecting the vari- ability found in this species in nature. Other entities in sect. Kneiffia have presented fewer taxonomic difficulties. Oenothera pilosella Raf. has been considered a variety of O. fruticosa in studies of Oenothera by Torrey & Gray (1840), Small (1896), and Léveillé (1902), but since Pennell’s monograph of Kneiffia, it consistently has been recognized as a distinct species. Oenothera sessilis (Pennell) Munz, likewise, was first recog- nized as a species by Pennell (1919), but in light of its close similarities to O. pilosella, both in overlapping morphological characteristics and like chromo- some number (n — 28), it is here assigned to subspecific rank within that species. The remaining three taxa, O. perennis L., O. spachiana Torr. & A. Gray, and O. linifolia Nutt. are morphologically distinct and have caused few taxonomic prob- lems in the past, other than a few inconsequential varieties having been described within these species from time to time. DiscussioN oF CHARACTERS Among the characters used in determining relationships among the species of Oenothera sect. Kneiffia and in evaluating the phylogeny of the section are the following: 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 385 Habit: Two species, O. linifolia and O. spachiana, are herbaceous annuals; whereas three, O. fruticosa, O. pilosella, and O. perennis, are perennial. A basal rosette of seedling leaves is produced in the annuals soon after germination. whereas an overwintering basal rosette is produced during late summer in the perennials. The rosette in the perennial species may be directly attached to the rootstock of the present years growth, forming a clump, or at the end of rhi- zomes several centimeters away from the current year’s rootstock, then forming a colony often covering several square meters. The rosettes remain evergreen through the winter and usually wither by early anthesis, although they may remain green during most of the flowering time. The primary stem begins to elongate in early spring as the photoperiod lengthens, only following a period of cold weather. In the perennials the underground base of the stem often becomes swollen and woody, especially in drier habitats. The formation of rhizomes is characteristic of O. pilosella subsp. pilosella, but rare in O. fruticosa subsp. fruti- cosa. A sparsely branched taproot is characteristic of the annuals. Fibrous roots (rarely fleshy in subxeric conditions) are characteristic of the perennial species. Stems are usually erect and are simple to much branched from near the base or above. When the stems become decumbent, they often root at the nodes. Leaves (Figs. 1-39): The leaves are variable in size, shape, pubescence, and texture. They range from linear or nearly filiform to ovate, with margins subentire to coarsely dentate, and often undulate. The basal leaves are always petiolate with attenuate bases, but the alternate cauline leaves usually become abruptly sessile or short petiolate. The leaves vary from thin and = translucent to quite thick and leathery, and are occasionally glaucous, especially beneath. The leaves are often dotted, streaked, blotched, or wholly red or purple. They are usually held at right angles to the stem or may be ascending. Pubescence: Most plants are pubescent throughout. Oenothera fruticosa subsp. glauca is the only taxon which is often nearly to quite glabrous through- out. In the other taxa stiff erect hairs are frequent near the base of the stem. Higher on the stems the pubescence also may be erect, as in O. pilosella subsp. pilosella, but it usually is incurved or appressed. The upper parts of the plants, especially the inflorescences, are usually more densely pubescent, with appressed or glandular hairs or a mixture of both. In O. linifolia, the pubescence of the inflorescence is puberulent or glandular-puberulent. The basal leaves are usu- ally glabrous except for their ciliate margins. The cauline leaves have less densely ciliate margins. Their surfaces are glabrous to densely strigose or occa- sionally with erect hairs, usually more pubescent above than below, and usually more densely so along the midribs. Hair color varies from whitish to grey or sometimes, especially in O. pilosella, tawny or yellowish. The pubescence varies under different growing conditions. During wet weather or in the relatively humid conditions of the greenhouse, glandular hairs become more frequent on the upper parts of the plant than on the same plant growing under drier condi- tions. In plants with mixed glandular and nonglandular hairs, the nonglandular ones become more frequent in more xeric conditions. Inflorescence: Distinct unbranched racemes or corymbs with few to many flowers occur in all but one species. The inflorescences may be either peduncu- 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 387 late and sharply distinct or not. The flowers are subtended by small bracts. The tip of the inflorescence in O. perennis and occasionally in O. fruticosa subsp. fruticosa nods and gradually becomes erect as the flowers open. In one annual species, O. spachiana, the flowers are borne singly in the axils of normal or slightly reduced cauline leaves on the upper two-thirds or half of the stems; here there appears to be a premium on flowering rapidly following germination. Flowers and Pollination Systems: The flowers of all species of sect. Kneiffia are pale to bright yellow, showy, and very faintly fragrant. Flowering begins in April in the southern portions of the distribution of the section and in June or July in the higher Appalachians and the northern United States and Canada, usually lasting a few weeks to two months, with occasional flowers until frost. The two annual species and one of the perennial species, O. perennis, are autog- amous, with the style about the same length as the filaments and pollen being shed directly on the enclosed stigmatic lobes often before the flowers open. The three remaining perennial species are allogamous, with the style longer than the filaments and the stigma held above the anthers. The sepals begin to reflex after dark the night before the flower opens and in the outcrossing perennials the stigmatic lobes often extend beyond the still tightly enfolded petals. The petals are probably too tightly folded together to allow any nocturnal pollinators to enter the flowers, even though the pollen usually dehisces during the night, and, at any rate, before the flower opens after sunrise the following morning. The flowers usually remain partially closed on cloudy days and close partially near sunset the first day. They may either wither that first night or reopen for at least six consecutive days before fading. The reflexed calyx often becomes flushed with anthocyanin and pinkish or purplish, as do the petals following pollination. The distal portion of the petals varies from cordate or emarginate to truncate and often undulate, either flat or + incurved. Cleistogamous flowers are frequently produced in O. spachiana and occasionally in O. linifolia and O. < FicunEs 1-39, Outlines of leaves in Oenothera sect. Kneiffia. Cauline leaves from about a third of the distance up the stem.—1-10. O. fruticosa subsp. fruticosa.—1. New Hanover Co., NC, Bell 13047 (VDB).—2. Monroe Co., WV, Straley 713.—3. Nottoway Co., VA Straley 1072.—4. Sussex Co., VA, Straley 705.—5. Faulkner Co., AR, branches tend to be few and are held at wide angles. In most other areas the branches are much more erect 405 406 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 2. Leaves: These are extremely variable in size and shape (Figs. 11-20). In plants from the higher Appalachians, the leaves are usually broadly elliptic to broadly ovate, becoming more lanceolate to very narrowly elliptic northward and at lower elevations. Cauline leaves are usually sessile, but they may have petioles up to 6 cm long as in collections from Amelia Co., VA, Straley 1078. Leaf margins are usually remotely dentate to subentire. They are typically more dentate than in most populations of O. fruticosa subsp. fruticosa. Among the collections with coarsely dentate leaves are those from Caldwell Co., NC, Ahles 43995 & Duke (NCU) and Rockland Co., NY, Lehr 886 (NY). Only at the higher elevations of the southern Appalachians do populations have leaves which are quite glaucous, especially beneath and particularly in the populations with the broadest leaves, which was clearly the reason for Michaux naming this taxon O. glauca. Many of the glaucous-leaved plants have very thick leathery leaves compared with those of plants from lower altitudes and higher latitudes. Among many examples of these thick, broad-leaved, glaucous collections are those from Patrick Co., VA, Kral 9262 (VDB) and Pike Co., KY, McVaugh 8695 (MICH, NA). Other broad-leaved collections are not at all glaucous and have very thin, translucent leaves as in collections from White House, KY, Biltmore Herb. 6739 (US) and Yancey Co., NC, Lloyd 4746 (MO). 3. Pubescence: Some plants are glabrous throughout, whereas others have strigose inflorescences and leaves, especially along the midrib and margins and stems. Most frequent are plants with short, erect, glandular hairs on the stems and especially the inflorescences. Glandular hairs usually predominate over nonglandular ones, and there are usually more glandular hairs on the young inflorescences early in the season. These apparently fall from the capsules as they mature. A few plants had long erect hairs (to 2 mm) on the stems and leaves, similar to those characteristic of O. pilosella subsp. pilosella: Watauga Co., NC, 22 June 1891, Small & Heller (F, MASS, PH, POM, YALE) and Cald- well Co., NC, 24 June 1893, Heller (MSC, PH). 4. Petals: These vary in size from 1 cm long in many plants from the north and at lower elevations to 2.5-3 cm long in those from the mountains. These larger flowers are frequently paler yellow than the smaller ones. Petals vary from emarginate to truncate and are often undulate distally. . Capsules (Figs. 42-43): These are the most variable structures in size, shape, and pubescence. At one extreme they are oblong and subsessile, and at the other extreme they are distinctly clavate and stalked, with all intermediate forms represented in different populations and many variations within popula- tions. They are usually widest near the middle, as measured from the base of the stipe to the distal end of the capsule body. The body varies from narrowly ridged to usually distinctly winged. Capsule shape often changes with maturity. Among some collections with very large and broadly winged capsules (23 mm long, 7 mm wide) are those from Patrick Co., VA, Kral 9262 (VDB There are many individuals and populations which are intermediate between the two subspecies of O. fruticosa. No single morphological character should be weighed too heavily in placing a plant in either of the subspecies. For example, nonglandular hairs have been regarded traditionally as a key characteristic of 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 407 O. fruticosa subsp. fruticosa, whereas, in fact, either subspecies may have glan- dular hairs, although they are more frequent in O. fruticosa subsp. glauca. In the southern Appalachians where O. fruticosa subsp. glauca is most distinct, there are few difficulties in separating it from subsp. fruticosa. In much of Tennessee, Kentucky, Ohio, Maryland, West Virginia, and Pennsylvania, how- ever, there are many intermediate populations and individuals, and the subspe- cific lines become much less distinct. By using a combination of characters, however, most specimens may be assigned to one or the other. In general, O. fruticosa subsp. glauca has broader, usually relatively glabrous, sometimes glau- cous, and more dentate leaves; predominately glandular pubescence; and oblong capsules, widest about the middle. Oenothera fruticosa subsp. fruticosa usually has narrower, more strigose leaves, with subentire margins; predominately non- glandular hairs; and clavate capsules, widest near the distal end. Among the populations which are intermediate between the two are: Lou- don Co., VA, Hunneywell 10718 (GH), with variable leaf width on different plants, and broadly winged capsules, widest at the middle, but all strigose, nonglandular pubescence; Jackson Co., OH, 22 June 1968, Bartley (OS), with clavate capsules, has two plants with glandular hairs and one with only non- glandular hairs; Grant Co., WV, 31 July 1931, Core (NY), with clavate capsules, but with only or predominately glandular pubescence. For states in which there has been an abundance of specimens of these two taxa collected, as in North Carolina, the pattern of distribution, overlap of char- acters, and intermediate populations is demonstrated quite well, with typical O. fruticosa subsp. fruticosa in the eastern part of the state at mostly lower elevations, and O. fruticosa subsp. glauca only in the western part of the state in the mountains. In other states, such as Kentucky, in which there has been much less collecting, the distribution and overlap of characters is less clear. These two taxa have not been found growing in close proximity, with the exception of one locality in Amelia Co., VA, Straley 1078, 1079 (MO), which was observed on 31 May 1975, where the two subspecies grow on opposite sides of a road, a few hundred meters apart. The population of O. fruticosa subsp. fruti- cosa was confined to a wet marshy area in the edge of a mixed deciduous woods, whereas that of O. fruticosa subsp. glauca was in the edge of a dry open meadow. The two were close enough for potential crossing by visiting bees but only the last flowers of subsp. fruticosa and the first flowers of subsp. glauca were open. At least in these populations, the possibilities of crossing were limited to overlap in flowering time of only a few days. No intermediates were found. Neverthe- less, the existence of many intermediate populations elsewhere makes it quite clear that hybridization between these entities is and has been very frequent and that the pattern of intergradation between them is complex and nearly complete. This taxon has been reported as tetraploid by Schwemmle (1924) and from plants cultivated at Edinburgh Botanical Garden by Hagen (1950). The origi- nal source of these collections is not known and no voucher specimens were made. The only other published reports of chromosome number for this taxon refer to O. fruticosa subsp. fruticosa, as already mentioned in the discussion of 408 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 that subspecies. A specimen from Clark Co., Virginia, Baldwin 5189 (GH), was annotated as a tetraploid by Earlene Atchison. Vouchers for chromosome number (34 ap ciet 12 5 Tetraploid, n = 14. New Jersey: Passaic Co., rh ale 760, 1114. NonrH CAROLINA: Mitchell Co., Lloyd 4744, 4 rings of 4, 1 chain of 4, 4 pairs. Transylvania Co.. Lloyd 4749. “wasa Co., Lloyd 4745. VIRGINIA: Amelia Co., of: 1078. Bedford Co., Straley 875. Craig Co., Straley 716, 720, 3 rings of 4, 2 chains of 4, 6 pairs. WEST VIRGINIA: vacan Co., Straley 756, 758, 5 rings of 4, 4 pairs. CULTIVATED: CANADA: NOVA SCOTIA: 1 Co., Straley 1103. UNITED STATES: PENNSYLVANIA: Chester Co., 5 ae 9 84 PA 2. Oenothera pilosella Raf., Ann. Nat. 1: 15. 1820. Perennial herb, + rhizomatous, from a thickened rootstock. Stem erect, simple or a few spreading to ascending branches above, 2-8 dm tall, densely to sparsely hirsute, the hairs 1-2 mm long, or strigose, the hairs less than 1 mm long on the stems, leaves, especially along the midrib and around the margins, and on the inflorescences, rarely glabrous. Basal leaves oblanceolate to ovate, 4-8 cm long, 2-5 cm wide, the petiole (0.5-)1-3(-4) cm long, entire, withering before flowering; cauline leaves mostly lanceolate, less commonly linear to ovate, 2-10(-13) cm long, 1-2(-4) cm wide, the petiole 0.1-0.5(-2) cm long, subentire to coarsely dentate. Inflorescences terminal, erect, and indistinct. Ovary (45-)8-12(-14) mm long, (1-)1.5-2.5(-3.5) mm thick, glabrous to sparsely pubescent with spreading hairs, or strigose, oblong or narrowly oblong, sessile or borne on a stipe 1-2 mm long. Floral tube 1-2.5 cm long. Sepals 10- 20 mm long, 2-5 mm wide, with free tips 1-3 mm long, connivent to divergent, + hirsute or strigose. Petals 1.5-3 cm long, 1.5-2.5 cm wide, obcordate to cleft, dark yellow. Filaments 7-15 mm long; anthers 4-8 mm long; pollen 145-165 um in diameter. Style 10-20 mm long; stigma held well above the stamens at anthesis; stigmatic lobes linear, 2-5 mm long, divergent. Capsule subsessile or stalked, the stipe (1-)3-5(-9) mm long, linear-clavate to elliptic, glabrous to densely hirsute or strigose, (7-)10-15(-20) mm long, 2-4(-5) mm thick, tetrag- onal, occasionally winged. Seeds dark reddish brown, ca. 1 mm long, ca. 0.5 mm wide, papillose. Gametic chromosome number, n — 28. Self-incompatible. Distribution ( Fig. 80): Open fields, edges of woods, and prairies, usually in somewhat marshy places from southern Ontario to Iowa, southward to northern Alabama and eastern Texas. Frequently cultivated and often escaping and be- coming established outside its natural range. Oenothera pilosella is a variable taxon which has been in the past frequently confused with O. fruticosa, but the two are actually sharply distinct. Some individuals or populations have individual morphological characters typical of the other taxon, but when overall morphology is considered, specimens are easily referred to one species or the other. This taxon consists of two subspecies. The first, O. pilosella subsp. pilosella, has a relatively broad distribution and is especially common in the Midwest and Mississippi River Valley. The second, O. pilosella subsp. demareei (O. ses- silis auct.) has, at present, a very narrow distribution in eastern Arkansas, central Louisiana, and the Gulf Coast of Texas. 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 409 Based on overlapping morphological characters, it seems best to regard these two entities as subspecies of a single species. They are exclusively octoploid (n = 28), and are the only octoploids known in the genus Oenothera. 2a. Oenothera pilosella Raf. subsp. pilosella. O. fruticosa L. var. hirsuta Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1: 496. 1840. LECTOTYPE: Illinois (NY). O. ee L. var. pilosella (Raf.) Small & Heller, Mem. Torrey Bot. Club 3: 26. 1892. Kneiffia fruticosa L. var. pilosella ( Raf.) Britton, X x Torrey Bot. Club 5: 234. 1894. K. pilosella ( Raf.) Heller, Cat. N. Amer. Pl., ed. 2 900. Migne roe L. £. hirsuta (Nutt. ex Torr. & x Gray) H. Lév., Monogr. Onoth. 108. Kneiffü sumstines O. Jennings, Ann. Carnegie Mus. 3: 480. 1903. TYPE: Pennsylvania, Arm- 5 pt Kittanning, upland field, June 1905, D. Sumstine (CM, holotype, not otype). K. pratensis de Fl. S.E. U.S, 842, 1335. 1903. rype: Missouri, Jefferson Co., wet places, 11 June 1878, H. Eggert I holotype: WIS, isotype). Oenothera ae (Small) B. L. Robinson, Rhodora 10: 34. 1908. O. pilosella Raf. f. laevigata Palmer & Steyermark, Brittonia 10: 116. 1958. rype: Missouri, Howell Co., 4 mi south of West Plains, J. Steyermark 78703 (F- 1456282, holotype; GH, MO, isotypes ). Rhizomatous perennial herb, from a thickened base, simple or little branched above, the branches spreading or ascending. Stems 2-8 dm tall, densely to sparsely pubescent with spreading hairs 1-2 mm long throughout, rarely gla- brous. Basal leaves oblanceolate to ovate, 4-8 cm long, 2-5 cm wide, the petiole (0.5-)1-3(-4) cm long; cauline leaves lanceolate to ovate, subentire to coarsely dentate, lanceolate to ovate, 3-10(-13) cm long, 1-2(-4) cm wide, abruptly narrowed to the base, subsessile or with a petiole to 0.5(-2) cm long. Ovary (4.5-)9-12(-14) mm long, (1-)1.5-2.5(-3.5) mm wide, glabrous or sparsely to densely pubescent with spreading straight hairs 1-2 mm long, narrowly ob- long, sessile or with a stipe 1-2 mm long. Floral tube 1-2.5 cm long. Sepals 10-20 mm long, 2-5 mm wide, the tips 1-3 mm long, usually divergent, hirsute. Petals 1.5-3 cm long, 1.5-2.5 cm wide, obcordate to cleft, dark yellow. Filaments 7-15 mm long; anthers 4-8 mm long; pollen 165 um in diameter. Style 10-20 mm long; stigmatic lobes linear, 2-5 mm long. Capsule linear-clavate to linear- elliptic (rarely elliptic), (5-)10-15(-28) mm long, 2-4(-5) mm wide, 4-angled to slightly winged. Gametic chromosome number, n — 28. Self-incompatible. Type: Indiana, Vanderburgh Co., near Evansville. Apparently no material of Rafinesque's original gathering has survived. Distribution (Fig. 80): Isolated colonies in wet meadows, edges of woods and prairies, southern Ontario to southern Wisconsin and southeastern Iowa to northern Alabama and central Louisiana. It is widespread in cultivation in gar- dens and frequently escapes and becomes naturalized; consequently, the north- ern and eastern natural limits of this taxon are not clear. Apparently, Wayne Co., West Virginia, along the Ohio River and Erie Co., New York, are the eastern limits and Tuscola Co., Michigan, and Mantiwoc Co., Wisconsin, the northern natural limits. Collections from the Upper Peninsula of Michigan are probably from naturalized populations, although they might represent natural disjunct 410 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 300 400 500 MILES Ficure 80. Distribution of Oenothera pilosella subsp. pilosella (circles) and subsp. sessilis (triangles). Dotted lines show the eastern and northern limits of the natural distri- bution. populations. It may be assumed that any populations in the Atlantic Coast states and Canadian Maritime Provinces are local escapes from cultivation. Illustration: Gleason (1963: 595). This is again a variable taxon, although not nearly so much so as O. fruti- cosa. Among the notable morphological characters which show considerable variation are the following: 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 411 1. Habit: Usually strongly rhizomatous, forming large colonies. Stems of many populations are all simple, others all branched toward the upper portions of the plant, or both simple and branched stems may occur within a single popu- lation. The branches are usually at wide angles to the main axis, or in some populations more strongly ascending. 2. Pubescence: The taxon is characterized by long (ca. 2 mm) erect hairs on the stems, inflorescences, and sometimes the leaves. The pubescence varies from white to yellowish or tawny. There are frequently subglabrous individuals within typical pubescent populations as in Jackson Co., OH, 29 June 1932, Bart- ley & Pontius (OS) and Athens Co., OH, Straley 980 (MO). The leaves are usually sparsely to densely strigose with shorter (1 mm) or longer (2 mm) hairs, but they may be subglabrous with ciliate margins. The mature capsules vary from densely strigose with hairs 2-3 mm long as in Monroe Co., MO, Hudson 518 (MO) to very sparsely pubescent or glabrous as in Howell Co., MO, Steyermark 78703 (F, GH, MO), the type of O. pilosella f. laevigata Palmer & Steyermark. 3. Leaves (Figs. 21-28): These are quite variable in size, shape, and pubes- cence. Cauline leaves are usually subsessile although they may be borne on petioles to 2 cm long. Leaf shape ranges from linear as in collections from Effingham Co., IL, Evers 16959 (ILLS) and Little River Co., AR, Tucker 16146 (MO), to elliptic or ovate as in Gallia Co., OH, Straley 925 (MO) and Knox Co., MA, Straley 787 (MO). Populations from the southern portion of the range tend to have narrower leaves than those from the north, but there are many exceptions and variations in leaf width within populations. Margins of leaves are subentire to remotely dentate. Leaves are usually held at nearly right angles to the primary axis, although they may be more or less ascending. 4. Capsules: "These vary in size, stipe length, shape, and pubescence. Size ranges from very short capsules 5-7 mm long which are sessile or on a short stipe (1 mm) as in Pope Co., AR, Tucker 15500 (MO), to long capsules 16-18 mm long on a stipe 10-12 mm long as in Adams Co., OH, July 1972, Bartley 23 (OS). Capsule shape varies from linear-elliptic or elliptic to linear-clavate. Among populations assigned to this taxon which approach O. pilosella subsp. sessilis in leaf shape and width and in capsule shape, but with long erect hairs on the stems, and with rhizomes are collections from La Salle Parish, LA, McVaugh 8482 (NA); Rapides Parish, LA, Kral 20065 (VDB); and Lonoke Co., AR, Demaree 22938 (BH, CU, ISC, MO, NO, NY, OKLA, TENN). There are no previous chromosome reports for this taxon except a probable octoploid (n= 28) (DeTurck, 1969). An octoploid chromosome number re- ported by Hecht (1942) from Cook Co., Illinois, as O. fruticosa doubtlessly refers to O. pilosella (A. Hecht, pers. comm.). Oenothera fruticosa does not occur near Chicago, whereas O. pilosella is (or was) frequent. Vouchers for chromosome number (24 individuals, 14 populations): Octoploid, n — 28. ARKANSAS: Ashley Co., Straley 1057, 1 ring of 8, 5 rings of 6, 1 ring of 4, 7 pairs. Cleveland Co., Straley 1069. Drew Co., Straley 1052. Omio: Athens Co., Straley 923, 933, 980. Gallia Co., Straley 925, 926, 927. Pike Co., Straley 943. CurrivATED: Kentucky: Harlan Co., Straley 694. Mane: Washington Co., Straley 802. Omio: Athens Co., Straley 924. Vir- GINIA: Giles Co., Straley 709. 412 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 2b. Oenothera pilosella subsp. sessilis (Pennell) Straley, comb. nov. Kneiffia sessilis Pennell, Bull. Torrey Bot. Club 46: 366. 1919. Oenothera sessilis (Pennell) Munz, Bull. Torrey Bot. Club 64: 291. 1937. Perennial herb 3-6.5 dm tall, from a distinctly thickened, + bulbous rootstock to 1 cm thick. Stems simple to few branched in the upper third, the branches ascending, densely strigose with hairs less than 1 mm long throughout, except near the base, where subglabrous. Basal leaves oblanceolate, subglabrous with ciliate, undulate margins, 2.5-7 cm long; 0.7-2.3 cm wide, the petiole 1-1.5 cm long; cauline leaves + ascending, lanceolate to narrowly lanceolate (3-)6-7(-9) em long, (0.3-)0.6-0.8(-1.1) em wide, sessile, subentire. Ovary 4.5-6.5(-8) mm long, 1-1.5(-2) mm thick, oblong, sessile. Floral tube 10-15(-20) mm long. Sepals 10-18 mm long, 2-3 mm wide, the tips 1-2 mm long, connivent to = divergent. Petals 1.5-2.5 cm long, 1.8-2.2 cm wide, obcordate, dark yellow. Filaments 7-9 mm long; anthers 5-8 mm long; pollen ca. 145 um in diameter. Style 10-12 mm long; stigmatic lobes 2-4 mm long. Capsule elliptic, 8-10 mm long, 3-4 mm thick, 4-angled, not winged, sessile or with a stipe 1-2 mm long. Gametic chromosome number, n — 28. Self-incompatible. Type: Arkansas, Little Rock Co., Little Rock, 2 June 1885, H. E. Hasse (NY). Distribution (Fig. 80): Presently limited to remnant wet or usually dry prairies of eastern Arkansas, central Louisiana, and the Gulf Coast of Texas, pos- sibly surviving only in eastern Arkansas. Illustration: Fig. 81. Specimens examined: ARKANSAS: Arkansas Co.: DeWitt, riceland prairies, Demaree 21078 (BH, MO, NY, SMU). Near Hagler, flat upland, Chamberlain 33 (ILL). 3 mi SW of Stuttgart, prairie, Straley 1071. Ashley Co.: Mist, prairies, Demaree 15086 (POM, SMU). Prairie Co.: Wheeler 57 (F, MICH). Near Hazen, Grand Prairie, Palmer 25038 (MO). St. Francis Co.: Forrest City, valley land, Demaree 15107 (BH, SMU). LOUISIANA: Tensus Parish: Hale s.n. (MICH). Texas: Galveston Co.: Galveston Island, Lindheimer s.n. (US). Wrruour Locaity: Hale s.n. (MASS, NY, P). Wrrnour Data: (NY, US). This taxon, named by Pennell (1919) and retained as a species by Munz (1937, 1965), is treated as a subspecies of O. pilosella, based on chromosome number and morphological similarities. It is rarely collected and differs from O. pilosella subsp. pilosella in the following characteristics. It is apparently not rhizomatous, although some populations may be so. In the only two extant populations which have been located, Arkansas Co., AR, Straley 1071 (MO) and Prairie Co., AR, Straley 1049 (MO), basal rosettes are produced directly from the rootstock of the current year or on a very short, + woody extension of the spherical root- stock which cannot be called a rhizome. Both the branches and leaves are more ascending than in O. pilosella subsp. pilosella and the leaves are typically narrower. The pubescence is of short (1 mm or less), dense, appressed hairs throughout. The sepal tips are usually + connivent in bud. The ovaries and mature capsules are usually elliptic with a short stipe. Collections from Ashley Co., AR, Demaree 15086 (POM, SMU) assigned to this taxon are somewhat intermediate to subsp. pilosella in the long erect hairs on their stems, but have the general aspect of O. pilosella subsp. sessilis. Another collection which 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 413 Ficures 81-82.—81. Oenothera pilosella subsp. sessilis (»Y4). [After Straley 1049 (MO).]—82. O. spachiana (x%). [After Emig 582 (N ] seems best assigned to subsp. sessilis is from Little Rock, AR, 26 May 1885 Hasse (NY) and is densely, short strigose throughout, lacking any long hairs. However, it has broader leaves, more typical of subsp. pilosella. A specimen with leaves of similar size and shape from near Corning, Clay Co., AR, 25 May 1893, Eggert (MO), has the strigose vestiture of subsp. sessilis although distinctly yellow- ish—more typical of subsp. pilosella. These collections lack good underground parts and mature fruits, with which they could be more definitely assigned to one subspecies or the other. Pennell’s type specimen of this taxon was presumably collected by Hasse in the vicinity of Little Rock, although the hilly uplands around the city of present- day Little Rock are somewhat out of line with the ecology of the presently known 414 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 populations of this subspecies. However, Hasse's collection may have been from the flatter, lowlands east or south of the city, or even many miles from the city. Munz was apparently confused between this entity and some populations of O. fruticosa from the same general area. Munz (1937: 291) states that O. sessilis is distinct “from O. fruticosa in the narrower, more elongate capsules," when, in fact, all of the mature capsules of this taxon are neither as narrow nor as elon- gated as capsules of O. fruticosa. However, most of the specimens of subsp. ses- silis which Munz probably saw have immature capsules. At any rate, the type collection, other specimens cited above, and the two known extant populations certainly represent a distinctive, yet apparently rare taxon. Vouchers for chromosome number (3 individuals, 2 populations): Octoploid, n — 28. ARKANSAS: Arkansas Co., Straley 1071. Prairie Co., Straley 1049. 3. Oenothera perennis L., Syst. Nat., ed. 10. 998. 1758. O. pumila L., Sp. Pl., ed. 2. 493. 1762, illeg. subst. Based on O. perennis L. O. pusilla Michx., Fl. Bor. Amer. 1: 225. 1803. TYPE: C nada, Québec, Lake Mistassini, in stony places, A. Michaux (P); Munz, Bull. Torrey Bot. Club 64: 303. 1937 O. chrysantha Michx., Fl. Bor. Amer. 1: 225. . TYPE: Canada, Québec, A. Michaux (P); Munz, Bull. Torrey Bot. Club 64: 303. 1937. O. pumila L. var. minima Lehm. in Hook., Fl. Bor. Amer. 1: 212. 1833. Based on O. pusilla Michx. Kneiffia pumila (L.) Spach, Hist Nat. Vég. Phan. 4: 377. 1835. K. chrysantha ( Michx.) Spach, Nouv. Ann. Mus. Hist. Nat. 4: 368. 1835. K. michauxii Spach, Ann. Sci. Nat. Bot., sér. 2, 4: 167. 1835, illeg. subst. Based on Oenothera umila L. and O. gracilis. Oenothera pumila L. var. pusilla (Michx.) Walp., Repert. Bot. Syst. 2: 84. 1843. O. pumila L. var. chrysantha Gordinier & Howe, Fl. Renssalaer Co., N.Y. 14. 1894. TYPE: New York, Renssalaer Co., Postenkill, E. C. Howe (not located ). O. pumila L. var. rectipilis S. F. Blake, Rhodora 19: 110. 1917. TYPE: Canada, New Bruns- wick, Petit Rocher, 21 Aug. 1913, Blake 5513 (GH, holotype; CU, NY, P, TEX, US, isotypes ). Kneiffia perennis (L.) Pennell, Bull. Torrey Bot. Club 46: 372. 1919. K. depauperata O. Jennings, J. Wash. Acad. Sci. 10: 454. 0. TYPE: Canada, Ontario, northeast of Sioux Lookout, shore of a boulder-strewn bay of the lake, 7 Sep. 1914, O. E. & G. K. Jennings 7501 (CM). Oenothera perennis L. var. rectipilis (S. F. Blake) S. F. Blake, Rhodora 25: 47. 1923. Perennial herb from fibrous rootstock. Stems simple or clumped and branch- ing above, usually erect to slightly decumbent, (0.3-)1.5-3(-7.5) dm tall, with short (ca. 0.5 mm long) straight or incurved hairs, the upper parts, especially the inflorescences, glandular puberulent. Overwintering basal rosette withered by early anthesis. Basal leaves oblanceolate to obovate, 2-4 cm long, 0.2-1.2 cm wide, glabrous except for the ciliate margins, the petiole (0.2-)0.5-1.2(-2.5) cm long; cauline leaves oblanceolate to obovate, 3-7 em long, 0.2-1.2 cm wide, narrowing to a winged petiole 0.1-1 cm long, sparsely strigose, the margins ciliate. Inflorescence nodding, glandular pubescent, relatively few flowered; subtending leafy bracts 8-18 mm long, 1-2 mm wide. Ovary 6-12 mm long, 1-2 mm thick. Floral tube 3-10 mm long. Sepals 2-4 mm long, 0.5-1 mm wide, the tips less than 1 mm long, connivent. Petals 5-10 mm long, 4-10 mm wide, trun- cate to cleft, dark yellow. Filaments 3-4 mm long, erect; anthers 1-2 mm long, shedding pollen directly on the stigma at anthesis; pollen ca. 108 Am across, 32-54% empty. Style 3-4 mm long, erect; stigmatic lobes about 1 mm long, di- 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 415 vergent. Capsule tetragonal or narrowly winged, clavate, glandular-puberulent, 2-10 mm long, 2-3 mm thick, tapering to a short stipe 1-2 mm long. Seeds bright rusty brown, 0.7-0.8 mm long, 0.2-0.3 mm thick, papillose. Gametic chromosome number, n= 7 (ring of 14 at meiotic metaphase I). Autogamous, often in bud; complex heterozygote. Lectotype: Miller, Figs. Pl. 2: 188. 1757. Drawn from a plant grown at Chelsea Physic Garden from seeds received from the Trianon, originally of Canadian origin. Distribution (Fig. 83): Fields, open woods, and boggy areas from New- foundland to southeastern Manitoba, south along the mountains to South Carolina. less common westward to Missouri. Isolated collections have been seen from coastal North Carolina, Hertford Co., (NCU) and British Columbia, New West- minster (CAN, UBC) and Shawnigan Lake, Victoria Island ( Melburn, 1965), where it is probably persistent as a garden escape. One collection, Macoun 482 (US) from "Saskatchewan Plains" is doubtfully from the province of Saskatche- wan, but may be from Saskatchewan Co., Manitoba. Illustration: Gleason (1963: 595). This is the most distinct and least variable taxon among the perennial species of sect. Kneiffia, but it does vary in the following characters: abit: Occasionally very short plants flower and fruit. An especially small collection only 3 cm tall is from Lake St. John Co., Québec, 24 July 1892, Kennedy (GH). Very tall plants (6.5-7.5 dm tall) are included in collections from Franklin Co., ME, Seymour & Potter 24333 (VT) and York Co., ME, Straley 1096 (MO). Stems are usually simple or few branched. A collection from Washington Co., ME, Fernald 1998 (NEBC) has 32 branches from the base. Others from Digby Co., Nova Scotia, Graves 22002 (PH) and Cheshire Co., NH, Massey & Boufford 4139 (NCU) are much branched higher on the plant. 2. Pubescence: Most plants have the lower two-thirds of the stem clothed with short (0.5 mm) appressed or incurved hairs. Some collections, particularly from eastern Canada, have stiff erect hairs (0.5 mm) on the stems and have been known in the past as var. rectipilis S. F. Blake. But single collections often include some plants with erect hairs and others with appressed hairs: Carleton Co., New Brunswick, 30 May 1952, Slipp (DAO) and Kings Co., New Bruns- wick, Straley 1098 (MO). Other collections—e.g., Montmorency Co., Québec, 28 June 1905, Macoun (GH, NY) and Missisquoi Co., Québec, Raymond Champagne 1401 (DAO)—include plants with either erect straight hairs, ap- pressed hairs, mixtures of these, or nearly glabrous stems. Occasional collections have densely strigose leaves clothed with white hairs to 0.5 mm long, among which are collections from Muskegon Co., MI, Bazuin 6597 (MICH) and Thun- der Bay District, Ontario, Garton 10468 (CAN ). 3. Leaves (Figs. 33-39): These differ in size, but there is little difference in shape compared with that in other perennial taxa of the section. Plants from the southern Appalachians often have narrower leaves. There are occasional col- lections with very narrow leaves (1.5-2 mm wide) as: Tompkins Co., NY, 15 416 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 .. - - - - = == — 2 =e — Ficure 83. Distribution of Oenothera perennis. June 1903, Jackson (WIS). An unusual collection from a bog in Muskegon Co., MI, Voss 9164 (MICH), is much branched, with widely spaced bractlike leaves (0.5-2 mm wide and 10 mm long) throughout most of the plant. 4. Inflorescences and flowers: The inflorescences are characteristically nod- ding in bud, gradually becoming erect as the flowers open. Frequently, de- pauperate plants with only a few flowers do not have nodding tips. Plants with unusually small flowers (petals 3-4 mm long) include: Hillsborough Co., NH, 12 July 1916, Batchelder (NEBC) and Guernsey Co., OH, Laughlin 1151 (OS). Larger-flowered collections (petals 10 mm or more long) include, among 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 417 others: Coos Co., NH, Pease 16403 (NEBC) and Hancock Co., ME, 14 June 1889, Rand & Renfield (NEBC). Plants from the southern part of the range often have smaller flowers. Cleistogamous flowers have not been observed in nature, but are suspected in collections from Grafton Co., NH, Fernald 11817 (NEBC) and Pon- tiac Co., Québec, Dore 20883 (TRT) in which part of the large, unopened flower buds are dropping from the ovaries, which otherwise appear normal. Examina- tion of the buds in these collections shows that pollen had been shed in the stigmatic lobes, as usual in the species. Valcanover (1927) reports a chromosome number of n = 14 for this taxon, with no reference to the origin of the population. In light of recent chromosome number determinations it is suspected that Valcanovers count actually refers to O. fruticosa. An herbarium voucher from Ottawa, Ontario, 12 July 1964, Mos- quin (DAO) was annotated as having a ring of 14 chromosomes by Raven. A report by Kapoor (1972) of 2n — 49 made from a root-tip preparation of a pop- ulation from Halifax Co., Nova Scotia, seems most doubtful. It seems virtually certain that Kapoor's 2n = 49 number refers to some other plant species and that his root-tip fixations or preparations were mixed. Kapoor (pers. comm.) also made a determination of n — 7 in plants of O. perennis from the same area but did not report it. This population was sampled in 1975 and only diploid plants with a ring of 14 chromosomes were found. Vouchers for chromosome number (59 individuals, 19 populations): n — 7. CANAD ADA: New Brunswick: Carleton Co., Straley 1113, ring of 14. Kings Co., Straley 803, 1098, ring of 14. York Co., Straley 1111, ring of 14. Nova Scorta: Halifax Co., Straley 1100, ring of SL ; Ki = C 1105. Queens Co., Straley 1107. UNITED STATES: MAINE: Knox Co., Straley 824, ring of 14. Waldo Co., Straley 793, 1097. York Co., Straley 1096, ring of 14. New HAMPSHIRE: Crafton ring of 14. Montgomery Co., Straley 753, ring of 14. West VIRGINIA: Nicolas Co., Straley 692. CuLTIVATED: Norway: Seeds from Botanical Garden, University of Bergen, Straley 693, ring of 14. 4. Oenothera spachiana Torr. & A. Gray, Fl. N. Amer. 1: 498. 1840. Blennoderma drummondii Spach, Nouv. Ann. Mus. Hist. Nat. 4: 407. 1835. LECTOTYPE: Texas, 1834, T. Drummond 81 (P; G, GH, K, isolectotypes ). Oenothera drummondii (Spach) Walp., Repert. Bot. Syst. 2: 85. 1843, non Hook., 1835. O. uncinata Scheele, Linnaea 21: 578. 1848. TYPE: Texas, Harris Co., prairie near Houston, Romer (not located ). Kneiffia spachiana (Torr. & A. Gray) Small, Bull. Torrey Bot. Club 23: 179. 1896. Onothera fruticosa L. race spachiana (Torr. & A. Gray) H. Lév., Monogr. Onoth. 106. 1902. Slender, erect annual herb 1-3(-4.5) dm tall from a sparsely branched taproot. Stems usually simple or with a few ascending branches from the base or higher, densely strigose throughout. Basal leaves 2-5 cm long, 0.5-1.5 cm wide, oblanceolate to elliptic, subentire, narrowing to a distinctly winged petiole 0.5-2.0 cm long, usually persistent to early flowering; cauline leaves narrowly oblong-lanceolate to oblong-linear, 3-6 cm long, 0.2-0.6 cm wide, the petiole 0.2-0.6(-1.5) cm long. Flowers in leaf axils of upper % or ? of plant. Ovary 8-12 mm long, 1-2 mm thick. Floral tube 4-10 mm long. Sepals 4-8 mm long, 1-2 mm wide, the free tips to 1 mm long. Petals 5-14 mm long, 10-15 mm wide, truncate to emarginate, pale yellow, turning pink, especially along the veins, 418 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficure 84. Distribution of Oenothera spachiana. after pollination. Filaments 3-7 mm long; anthers 2-4 mm long, shedding pollen directly on the stigma in bud; pollen ca. 70 um in diameter. Style 3-7 mm long; stigmatic lobes linear, connivent, 1-2 mm long. Fruit broadly clavate, 4-angled, 5-15 mm long, 3-5 mm wide, sessile, with a narrowed sterile base. Seeds straw colored, 1 mm long, 0.5 mm wide, verrucose. Gametic chromosome number, n = 7. Autogamous, usually in bud, often cleistogamous. Type: Texas, T. Drummond 81 (P, holotype; GH, isotype). Distribution (Fig. 84): Prairies, open roadsides and sandy places from eastern Oklahoma and eastern Texas to southeastern Arkansas and Louisiana. Populations in Panola and Lincoln counties, Mississippi and Hale Co., Alabama are apparently native, though possibly introduced. This species has also been collected as a ballast weed in Camden Co., New Jersey (NA). Scattered, but forming large populations. Illustration: Fig. 82. A relatively uniform taxon, morphologically, in pubescence, leaf and capsule size and shape, but variable in the following characters: Habit: Usually simple stems but may be much branched from the base (8-15 branches) as in collections from West Feliciana Parish, LA, Cooley & Brass 414 (GH, NCU) and Van Zandt Co., TX, Cory 57378 (SMU). Unusually small plants (7-9 cm tall) in flower are from Brazoria Co., TX, Palmer 5047 (POM). 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 419 2. Flouers: Collections with small flowers (petals 5-6 mm long) include those from Marshall Co., OK, Goodman 5823 (GH, OKL) and St. Augustine Co., TX, Palmer 9486 (US). Large-flowered collections ( petals to 15 mm long) include those from Little River Co., AR, Moore 510120 (WISC) and Grant Parish, LA, Shinners 31703 (OKL, SMU). Cleistogamous flowers are frequent in this taxon, with some populations apparently having all flowers cleistogamous. Some plants form both normal and cleistogamous flowers, and the proportion is probably under environmental control. Notable collections with cleistogamous flowers include those from Hampstead Co., AR, Bush 1428 (GH, MO, RSA) and Marshall Co., OK, Waterfall 11448 (OKLA). Plants raised from seed from Union Parish, LA, Straley 751 produced only normal flowers early in the season and only clesitogamous flowers later. The following year progeny from this collection produced plants with only cleistogamous flowers. Gregory & Klein (1960) report this taxon as diploid with 7 bivalents during meiosis from collections raised from seed from Marshall Co., OK, Waterfall 11448 (OKLA, RSA). Voucher for chromosome number (3 individuals, 1 population): Diploid, n — 7. Lovu- ISIANA: Union Parish, Straley 751 Subsection II. PENIOPHYLLUM Oenothera sect. Kneiffia subsect. Peniophyllum (Pennell) Straley, comb. nov. Based on Peniophyllum Pennell, Bull. Torrey Bot. Club 46: 373. 1919. Oenothera subgen. Kneiffia (Spach) mes sect. . Pe 5 (Pennell) Munz, Bull. Torrey ot. Club 64: 288. 1937; N. Amer. Fl., ser. 2, 1965. Erect annual herbs. Stems cue or with many ascending branches, villous near the base. Basal leaves ovate to obovate; cauline leaves linear. Inflorescences erect, strigulose to glandular puberulent; floral bracts shorter than the subtend- ing ovaries. Sepals without free tips. Petals pale yellow, flushed with pink after fertilization. Stigma surrounded by anthers at anthesis, the lobes blunt. Flowers sometimes cleistogamous. Capsules sessile, ellipsoid, rhomboid, 4-ridged. Ga- metic chromosome number, n = 7. Autogamous or cleistogamous. Type species: Oenothera linifolia Nutt 5. Oenothera linifolia Nutt., J. Acad. Nat. Sci. Philadelphia 2: 120. 1821. Kneiffia linifolia ( Nutt.) Spach, Nouv. Ann. Mus. Hist. Nat. 4: 368. 1835. K. linearifolia Spach, Ann. Sci. Nat. Bot., sér 2, 4: 167. 1835, illeg. subst. Based on Oenothera linifolia Nutt. Peniophyllum linifolium (Nutt.) Pennell, Bull. Torrey Bot. Club 46: 373. 9. Oenothera linifolia Nutt. var. glandulosa Munz, Bull. Torrey Bot. Club 64: 289. 1937. TYPE Georgia, DeKalb Co., Little Stone Mountain thin aad overlying rocks, 11 May 1901, A. Curtiss 6778 (GH, holotype; G, GA, GH, ILL, K, NA, NY, OKL, P, US, isotypes ). Erect annual herb 1-5 dm tall from a sparsely branched taproot. Stems sim- ple or with a few to many ascending branches from near the base or higher. Short erect hairs (0.2-0.4 mm long) near the base, occasional hairs on the margins of the leaves, the plants strigillose to glandular puberulent above, especially in the inflorescences. Basal leaves ovate to obovate or narrowly elliptic, 1-2(—4) em 420 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficure 85. Distribution of Oenothera linifolia. long, 0.2-0.6 cm wide, narrowing to a winged petiole 0.2-1(-1.5) cm long, sub- entire to remotely dentate, glabrous to sparsely strigillose or glandular puberu- lent, especially along the petiole, usually not persistent to flowering time; lower cauline leaves becoming abruptly sessile and linear or filiform, less than 1 mm wide, 1-4 cm long, crowded. Inflorescences terminal, distinct, unbranched spikes (1-)3-6(-12) cm long, glandular puberulent to strigillose, the hairs 0.1-0.2 mm long; bracts ovate to deltoid-ovate, 0.5-2 mm long, 1-3 mm wide. Ovary 3-6 mm long, 0.5-1.5 mm thick. Floral tube 1-2 mm long. Sepals 1.5-2 mm long, 0.3-0.6 mm wide, without free tips. Petals 3-5(-7) mm long, 1-3(-4) mm wide, obcordate to cleft, bright yellow. Filaments 1-2 mm long; anthers 0.5-1.0 mm long, shedding pollen directly on the stigma before and during anthesis; pollen ca. 93 jum in diameter. Style 1-2 mm long; stigma 0.5 mm long, shallowly 4-lobed. Capsules sessile (or with short stipe 1-4 mm long), 4-6(-10) mm long, 1.5-3 mm thick, ellipsoid-rhomboid, 4-ridged. Seeds pale reddish brown, ca. 1 mm long, ca. 0.5 mm wide, minutely verrucose. Gametic chromo- some number, n — 7. Autogamous, usually in bud; frequently cleistogamous. ype: Arkansas, summits of arid hills and the shelvings of rocks, near the banks of the Arkansas River, T. Nuttall (PH-910101, holotype; BM, GH, K, NY, PH, US, isotypes). Distribution (Fig. 85): Prairies, open rocky and sandy places, and open 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 421 roadsides, southern North Carolina to northern Florida, westward to southern Illinois, eastern Kansas and central Texas. Illustration: Gleason (1963: 595). This is the most distinctive and morphologically uniform taxon in the section. Variability in size and branching appears to be due largely to soil fertility and moisture levels during the growing season. Among the notable variations are the following: l. Habit: Although the plants are usually simple, they may be much branched from the base as in collections from Poinsett Co., AR, Demaree 29072 (SIU) and Johnston Co., OK, Correll & Correll 25018 (TEX), with 12-20 branches. Other collections exhibit much branching higher on the plant (30-50) branches, or ascending secondary branches as in a collection from Washington Co., AR, Wells 22 (US), with a thickened basal stem 5-6 mm in diameter near ground level, and in one from Independence Co., AR, Demaree 26936 (RSA). Branching appears frequently to be the result of mechanical injury to the apex of the plant. The basal seedling rosette of leaves sometimes remains green dur- ing early flowering, probably in response to adequate moisture levels in the soil, as in collections from Stone Co., AR, Demaree 64562 (SMU) and Latimer Co., OK, Hopkins 1685 (POM). 2. Pubescence: Some collections with predominately glandular hairs on the inflorescences have been known as O. linifolia var. glandulosa Munz. There are, however, many individuals and populations intermediate to the more fre- quent form with predominately nonglandular, incurved hairs in the inflorescence. The continued taxonomic recognition of this variety appears to be without merit. Flowers: Occasional individual plants or populations have very large flowers (petals 6.5-7 mm long) as in collections from Tangipahoa Parish, LA, 20 Apr. 1963, Wilson (FSU) and Drew Co., AR, Demaree 14578 (NY). 4. Capsules: The shape varies from subglobose and scarcely angled, with the body of the capsule ca. 4 mm long and the stripe ca. 1 mm long, as in collec- tions from Panola Co., TX, Shinners 20151 and Randolph Co., AL, McVaugh 8609 (MICH, TEX, US), to very elongated with the body ca. 6 mm long and the stipe 3-4 mm long as in collections from Izard Co., AR, Demaree 31792 (GH) and Payne Co., OK, 26 May 1916, Learn (OKLA). One previous report of chromosome number of n = 7, with 7 pairs, was made by Gregory & Klein (1960) from Montgomery Co., AR, Munz & Gregory 23503 (RSA). Vouchers for chromosome number (8 individuals, 4 populations): Diploid, n= 7. An- KANSAS: Conway Co., Straley 967, 7 pairs, 969, 7 pairs, 974. Prairie Co., Straley 961. LITERATURE CITED ALLEN, T. F. 1870. The Oenothera of Montauk Point, Long Island. Bull. Torrey Bot. Club 2 : .` Beeks, R. M. 1955. Improvements in the squash technique for plant chromosomes. Aliso BLAKE, S.F. 1918. Notes on the Clayton Herbarium. Rhodora 20: 21-28, 48-54, 65-73. CLELAND, R. E. 1972. Oenothera Cytogenetics and Evolution. Academic Press, New York. 492 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Dr ANDRADE, A. 1972. Oenothera x teframila > Rea 5: DeTurcx, J. E. 1969. A biosystematic study of selected Tp e di the genus Oenothera, ge 19 (Onagraceae). Ph. D. 5 Catholic Univ. of America, Wash- ington > DIETRICH, W. 1977. The cw a species of Oenothera sect. Oenothera ( Raiman- nia, Renneria; Onagraceae ). n. Missouri Gard, 64: 425-626. FERNALD, M. L. 1950. Gray's d of Botan . 8. American Book Co., New York. GLEAsON, H. A. 1963. The New Britton and ion Illustrated Flora of the Northeastern United States and Adjacent Canada. New York Botanical Garden ork. Grecory, D. P. & W. M. KLEIN. 1960. Investigations of meiotic t of six genera in the hace Aliso 4: 502-520. HacEN, C. W., 1950. A contribution to the cytogenetics E a genus Oenothera with special AUN to certain forms from South America. In R. Cleland (editor), Stud- ies in Oenothera cytogenetics and phylogeny. Indiana Univ. Eu Sci. Ser. 16: 305- Hecut, A. 1942. Colchicine-induced tetraploidy in Oenothera. Proc. Indiana Acad. Sci. 51: 41-44. 1950. Cytogenetic studies of Oenothera, subgenus Raimannia. In R. E. Cleland pu Studies in Oenothera cytogenetics and phylogeny. Indiana Univ. Publ. Sci. Ser. 16: Hooker, W. 3. 1837. Oenothera fruticosa var. ambigua. Bot. Mag. 64: sub 1 3545. KAPOOR, B. M. 1972. IOPB chromosome number reports XXXV. Taxon 21: 162. Laws, H. E. 1963. Polyploidy in Oenothera fruticosa L. J. Elisha Mitchell Sci. Soc. 79: 4 — 965. Pollen- grain 5 of polyploid Oenothera. J. Heredity 56: 18-21. Mon L£vEILLÉ, 1 1902 nographie du Genre Onothera. Institut de “y saqey Le Mans. Lewis, H. & M. E. Lewis. 1 5 5 The genus Clarkia. Univ. Calif. Publ. Bot. 241-392. LiNpgn, R. 1954. Etude génétique des mécanisms qui limitent la fertilité phe fae missouriensis et Oenothera fruticosa. Année Biol. 30: 501-518. MELBuRN, M. 1965. Small sundrops. Victoria ie 22: 4-5. MICHAUX, A. 1803. Flori Boreali-americana. Paris Muwz, P. A. 1937. Studies in Onagraceae. X. The subgenus Kneiffia (genus Oenothera ) and miscellaneous new species of i cid Bull. Torrey Bot. Clu -306. 965. Onegraceae. N. Amer. Fl, ser. 2, 5: 1-278. New York Botanical Garden, New York. PENNELL, F. W. 1919. A brief B een of the species B Kneiffia with the characteriza- tion of a new allied genus. Bull. Torrey Bot. Club 46: 363-373. Purs, F. 1814. Flora Americae ee White, Cochrane & ondon Raprorp, A. E., H. E. Antes & C. R. Bett. 1968. Manual of the a Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill. Raven, P. H. 1964. The generic subdivision of Onagraceae, tribe Onagreae. Brittonia 16: 276-28 . P. Grecory. 1972a. Observations of meiotic chromosomes in Gaura. Brit- ae 24: 71-86. — — — & ————. 1972b. A revision of the genus Gaura (Onagraceae). Mem. Torrey Bot. Ch b 23: 1-96. SARGENT, C. S. 1889. Portions of the journal of André Michaux, botanist, written gue his travels in the United States and Canada 1785-1796. With an introduction and explana- tory notes by C. S. Sargent. Proc. Amer. Philos. Soc. 26: dud cedes J. 1924. Vergleichend zytologische Untersuchungen an Onagraceen. Ber. sch. Bot. Ges. 42: 238-243. atr. 1. K. 1896. Oenothera and its segregates. Bull. Torrey Bot. Club 23: 167- M Spacu, E. 1835. Monographia onagracearum. Nouv. Ann. Mus. Hist. Nat Tuwarres, R. G. 1904. Early western travels 1747-1846. III. Arthur H. Clark, ntl Torrey, J. & A. Gray. 1838-1840. Flora of North America. Wiley & Putnam, New rk. VALCANOvER, R. 1927. Contributions a l'étude de la réduction ERU Oenothera biennis. Cellule 37: 203-229. WarsoN, S. 1873. Revision of the extra-tropical North American species of the genus Oenothera. Proc. Amer. Acad. Arts 8: 573-618. 1977] STRALEY—OENOTHERA SECT. KNEIFFIA 493 INDEX TO SCIENTIFIC NAMES Numbers in boldface type refer to descriptions; numbers in roman type refer to synonyms; numbers with daggers (+) refer to names incidentally mentioned. Blennoderma 396 drummondii 3967, 417 Bombus 3887 aura coccinea 3947 Halictidae 388+ Kneiffia 3827, 3847, 394 alleni 397 angustifolia 397 7 fruticosa 397 var. differta 403 —var. humifusa 398 —var. pilosella 409 š — 5 ag 2 2 ac — = m". mu 2 ce TN 3951, 3967, latifolia 403 linearifolia 419 Husa pi -1 var. b ida var. an 398 veluti Oenothera ES 3847, 4097 . Kneiffia 394, 3967 3 Eukneiffia 3967 —sect. Kneiffia 396 —sect. Peniophyllum 419 —sect. Blennoderma 396 —sect. Kneiffia 3817, 3827, 3837, 384f, 3877, 3887, 3891, 3907, 3917, 3927, 3937, 394, 3947, 396, 4157 —subsect. Eukneiffia 3817, 3937, 3947, 396 —subsect. Peniophyllum 3817, 393+, 47, 419 . Oenothera 3827 40: drummondii 417 florida 397 fraseri 403 fruticosa 381, 3827, 383t, 3847, 385%, 897, 391t, 3937, 3947, 3957, 396, 3977, 4067, 4081, 4107, 4117, 4147, TT —subsp. — ih 3831, 3851, 3871, 3907, 3927, 3937, 3941, 3951, Add 19 5 4007, 4017, 4021, 4047, 4057, 406, —subsp. glauca Xa 3837, 3857, 3871, „3897, 3907, 3927, 3937, 3941, 3951, 396t, 3971, 3997, 4017, 4027, 403, 4057, 4067, 407 ar. ambigua 403 — var. angustifolia 397 var. differta 403 — var. eamesii 398 — var. fraseri 403 —var. goodmanii 398 — var. hirsuta 409 —var. humifusa 397, 4007 — var. incan : — var. indic —var. opis 397 var. linearis E^ —var. pie 398 —var. vera glauca 3831, 5 . 4067 var. fraseri gracilis 4147 hybrida 403 — var. ambigua 403 403 var. eamesii 397, 400+ Tu 381t, 3847, 3857, 3871, 388+, 89t, 391, 3927, 3937, 3947, 3951, oe OF —var. glandulosa 419, 421+ longipedicellata 39 perennis 3811, 3831, 3847, 3857, 3871, 3887, 3897, 391t 3927, 393t, 3947, 4 1 : 157 pilosella pr 3841, 3857, 391+, 3947, 8, 4 424 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vor. 64 subsp. pilosella 381, 3857, 3877, — var. brevistipata 398, 4027 388, 3897, 390f, 3937, 3947, 395f, — var. fraseri 403, 4041 4017, 406+, 408t, 409, 4107, 4127, — var. fraseri 4 —f. hybrida 403 subsp. sessilis 3817, 3871, 3887, 3897, —f. latifolia 403 390t, 3927, 3937, 3947, 3957, 4087, — var. longistipata 398 4107, 4111, 412, 4137 var. riparia 398 —f. laevigata 409, 4117 —var. tetragona 402+ ratensis : var. velutina 398 pu uncinata 417 var. "waa 414 Onagraceae 381 —var. minima 4 ' nothera — var. pusilla 414 fruticosa var. rectipilis 414 —f. angustifolia 397 pusilla 414 —f. diversifolia 397 Manis 397, 400t —f. lucida 403 se a 403 —f, hirsuta 409 sae "3841, 4127, 4147 race spachiana 417 spachiana 3817, eg 3857, 3871, 3887, —var. angustifolia 397 3897, 3917, 3927, 3937, 3947, 3967, —var. glauca 403 , 418+ —var. maculata 403 subglobosa 398 Papilionidae 3887 —var. arenicola 398 Peniophyllum 419+ anne 3827, e gk 403 linifolium 419 subsp. glau Pieridae 388+ var. riparia xo Raimannia 3827 —subsp. tetragona Syrphidae 3881 — var. sharpii 39 THE SOUTH AMERICAN SPECIES OF OENOTHERA SECT. OENOTHERA (RAIMANNIA, RENNERIA; ONAGRACEAE '? WERNER DIETRICH? ABSTRACT s paper 5 an account of the native and naturalized South American species of 8 sect. Oenothera ( Onagraceae). Included are 63 taxa (species and subspecies) of Raimannia 3 (O. laciniata subsp. laciniata, O. laciniata subsp. pubescens, O. drummondii), and subsect. Euoenothera also 3 ( . erythrosepala, O. villosa, O. biennis e main focus of this monograph is a revision of the exclusively South American sub- sect. Munzia, which is divided into the three series Renneria, Allochroa, and Clelandia. All taxa were pa aoe at the Botanical Institute of the University of Düsseldorf from one to ' Dedicated to my wife and my children Nikola, Lorenz, Bernadette, Gabriel and Christoph. * I would like to thank those who have made the completion of this study possible. Prof. Wilfried Stubbe not only allowed me to work on this problem but also most generously pro- Raven made it possible for me to for six months at the Missouri Botanical Garden, during which time I was also able to visit South America for two months ork in St. Louis and in South America was su b grant from the U.S. National Science Foundation to pp n, who also translated this manuscript from German. His interest and cooperation were invaluable for my work, and useful discussions WEM. it were also held with Prof. Stubbe, Dr. A. Basler, Dr. M. Drillisch, and Dr. H. Kutzelni Thanks are due to Prof. K. and Dr. K. Santarius, E undertodk a five-month trip to South America in 1968 for the limosa of collecting sead samples of Oenothera and thereby added more than 2,000 strains to the collection at the University of Düsseldorf. Their trip, present study possible. Mr. J. D. Conra accompanied me to South America in January and February, 1974, and helped greatly to make our trip a success. Many South 1 bot- anists contributed in one way or another to the success of our trip, but I wou o thank especially Dr. E. Bordaz (Asunción, Paraguay), the late Prof. Dr. A. Burkart rte Isidro, Argentina), F. Encarnación (Lima, Peru), Prof. Dr. A. T. Hunziker (Córdoba, m. R. M. Klein (Itajai, Brazil), L. F. Lantensohlager ( Asunelón, Paraguay), and Prof. . E. Vanzolini (Sáo Paulo, Brazil). y technical assistant, Mrs. L. Mencke, tirelessly and expertly performed the necessary ud in the laboratory and garden that helped to make this study possible. I would also like to thank the gardeners of the University of Düsseldorf for their cultivation of Oenothera over a period of many years, and many students at the University who performed a variety of technical functions on these plants. Many botanists kindly contributed seeds for the study, and I am grateful to them. The University of Düsselfdorf and the Minister für Wissenschaft und Forschung des Landes Nordrheinwestfalen aw E me a “Forschungsfreisemester” for the 3 of this study, and I am most grateful to ther Finally, I would like to thank my wife and four children for their patience and under- standing. I hope that the successful conclusion of my work represented by this revision len help to make up in part for the six months that we were apart during the course of i completion. Material from the following herbaria was examined, and I am grateful to those who made the material in their charge available for study: AAU, AI (by Raven), AK, AMD, B, BA, BAA, BAB, BAS, BB, BM, BR, BREM, C, COI, CONC, CORD, CTES, DUSS, E, F, FR, C, GH, GOET, HB, HBG, HBR, K, L, `LD, LE, LIL, LISE, LISU, LP, LY, M, MEL, MICH, MO, MPU, MVFA, ND, NSW, NY, P, PERTH, POM, PR, PRC, PRE, R, RAW (by Raven), RB, xdi S, SGO, SI, SP, SRGH, UC, UPS, US, USM, W, Z, and the private Herbarium of D he Ruiz Leal, Godoy Cruz, Mendoza, Argentina T D. ° Botanisches Institut of the University of Düsseldorf, German ANN. Missouni Bor. GARD. 64: 425-626. 1977. 426 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 several wild strains. Cytological examination of meiosis in 600 plants representing 280 strains uar that 20 taxa were chromosomally homozygous, forming 7 bivalents at meiotic meta- phase I. All other taxa were complex heterozygotes forming a ring o chromosomes at meiotic metaphase I or with a smaller but still stable ring configuration (ring of bivalent, ring of 8 and ring of 6, or ° of 10 and ring of 4). Homozygous species distr ibuted throughout the range of subsect. 1 Some of the individual 5 homozygous species also had individual S Ida ranges; for example, O. scabra and O. versicolor series Renneria), O. mendocinensis, O. odorata, O. indecora, O. affinis, and O. ravenii (series Allochroa The following species of subsect. Munzia were adventive or established outside of South America: (1) O. mendocinensis; Europe (adventive); (2) O. longiflora subsp. longiflora: Europe, Atlantic Islands (Azores, Madeira, Canary Islands, Cape "ande Í wow South Africa, Australia; (3) O. indecora subsp. indecora: Europe ( adventive); (4) O. indecora sub m. bonariensis: Europe (adventive), Australia, South Africa, Tristan da Cunha (adventive?); (5) O. affinis: Europe, India, Pakistan, Australia, prie South Africa; (6) O. 5 U.S. adventive?), Australia; (7) O. stricta subsp. s ricta: U.S.A., Mexico, Hawaii, Europe, Japan, India, Pakistan, Sri Lanka, Java, Australia, M Zealand, North Africa, South Africa, Ethiopia; (8) O. parodiana subsp. 1 South Africa, Europe (adventive). The common ancestor of Oenothera sect. Oenothera subsect. unzia seems to have arriv ed i in South A America from North America no more than a few million years ago, and to Ee — c 2 Š = re c = c = = S Qz ~= a = S — a Š = ° > a wv — — Nn an ein elev aon of the mountains in 0 pubescens seems to have arrived in So uth America from North America much more recently, perhaps within s past 100,000 vears. Oenothera has been the subject of genetic and evolutionary investigations for a very long time, owing mainly to its unusual genetic system, which involves complex heterozygosity. The species of the so-called “Euoenothera” group of North America have been intensively studied for more than 70 years (summary, Cleland, 1972), but other members of the genus have not been considered in such depth. In Düsseldorf, a program has been conducted for more than 15 years on the biosystematics of the South American species of Renneria ( Fischer, 1962) and Raimannia (Munz, 1935). In this work an extraordinarily rich array of wild populations of the group assembled under the direction of Professor Doctor Wilfried Stubbe has been utilized. The object has in part been to work out a more satisfactory taxonomic rearrangement of the South American species of these two closely related groups. Extensive investigations of the genetics of both groups had been carried out earlier by Professor J. Schwemmle of Erlangen and his students, but the results have not, for the most part, been applied to the systematics of these taxa. ybridization has played an important role in the evolution of many groups of the genus Oenothera, so that a satisfactory taxonomy often can be worked out only on an experimental basis. In the course of my investigations with hundreds of strains of living plants at the University of Düsseldorf, I was able to ascertain that the most recent revision of subgenus Raimannia (Munz, 1935) was inade- quate in dealing with the rich pattern of variation in this group. It was soon evident that there were many more distinct entities than had previously been thought to exist; the phenomenon of complex heterozygosity had not been taken into account in earlier efforts to classify the subgenus. In the case of Renneria, no revision of the entire group has hitherto been available. In the more recent floristic and taxonomic literature all species have been treated under the names Oenothera campylocalyx and O. rubida. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 497 The common ancestor of what have been regarded as the subgenera Euoeno- thera, Raimannia, and Renneria of Oenothera almost certainly originated in the semiarid to subhumid regions of western North America in mid-Tertiary time (Raven, pers. comm.). This region is, in fact, the center of evolution of the entire tribe Onagreae, to which Oenothera belongs. Geologically speaking, the arrival of Oenothera in South America seems to have been very recent, an event of the past several million years (Raven & Axelrod, 1974). Aside from five of the ten species of Hartmannia (Raven & Parnell, 1977) and two of Lavauxia, all indigenous South American species of Oenothera are included in sect. Oenothera and are treated in this revision. Considering the reticulate relationships between the species grouped by Munz (1965) as the subgenera Oenothera and Raimannia, as well as the com- plex pattern of relationships among the South American species of this group, it seems preferable to regard all as belonging to a single infrageneric taxon. This taxon is well differentiated from all other groups in the genus, both morpholog- ically and in terms of its crossability. In accordance with the practice estab- lished for all other genera of Onagraceae, this group is designated a section, sect. Oenothera. 'The South American species of this section, all but four of which are assigned to the newly described, endemic subsection Munzia, are the subject of this revision. Oenothera sect. Oenothera subsect. Munzia is regarded as comprising three series and 45 species, of which three are divided into two subspecies each and five others are divided into three subspecies each, for a total of 57 taxa. One or more strains of each of these were cultivated in the experimental garden at the Botanical Institute of the University of Düsseldorf. All plants examined cytolog- ically were diploid, 2n — 14. The cytological examination of 600 plants repre- senting 280 strains showed that 20 species formed pairs of chromosomes at mei- otic metaphase I. All others were complex heterozygotes that formed either a complete ring of 14 chromosomes at meiotic metaphase I or a stable ring configu- ration (ring of 12 and 1 bivalent; ring of 8 and ring of 6, or ring of 10 and ring of 4) Many of the species of subsect. Munzia vary in such a way that they could be subdivided into several more species if the pattern of distribution of their morphological characteristics were taken as preeminent. In other words, if all of the small differences within the subsection that are preserved by the syndrome of self-pollination and complex heterozygosity, and which have in many cases come to characterize populations, were to be recognized taxonomically, one could recognize literally hundreds of species without any gain whatsoever in taxonomic utility. ere are many direct genetic connections between the South American species recently referred to subg. Renneria (Fischer, 1962) or earlier to subg. Euoenothera (Munz, 1933) and those referred to subg. Raimannia (Munz, 1935). The South American species referred to these two groups are completely interfertile with one another. Judging by the patterns observed, hybridization seems often to have occurred in nature, and at least a dozen species, referred in this monograph to series Clelandia, seem to have originated as complex 428 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 heterozygotes between these two morphologically quite distinctive groups. The species assigned to this series combine one genome derived from series Renneria with another derived from series Allochroa. The group is not, however, to be re- garded as a transitional one between these two series; no such transition exists. Rather it has originated as a result of hybridization between individual species of Renneria and individual species of Allochroa, as a result of which the species share a number of characteristics in common. The differentiation of the narrow- fruited ancestors of series Allochroa took place first, evidently from plants that resembled those now assigned to series Renneria. Presumably this took place at the southern end of the area of distribution of series Renneria, which is the region of overlap between the two series at the present time ( Fig. 7), and there- fore the area in which the species of series Clelandia are concentrated. It ap- pears likely that many of the species of this series might have had a very recent origin; with the exception of O. magellanica and O. punae, each has a restricted area of distribution. Genetic relationships thus seem to demonstrate a unity between South Ameri- can species of Raimannia and the group that has recently been called Renneria. On the other hand, the North American species of Raimannia, including O. laciniata, do not appear to be closely related to any species of this South Ameri- can group as its members are to one another, and they are here regarded as comprising a separate subsection Raimannia. This group is native only in North America with the exception of the entity that has been called O. laciniata subsp. pubescens, which ranges south to Colombia, Ecuador, and Peru. To series Renneria (13 species) belong all species with short, urn-shaped capsules, except for one element within the species Oenothera nana which was put in sect. Raimannia by Munz (1935); see p. 488. Series Allochroa (21 spe- cies) comprises only a portion of the South American species of Raimannia, for which a new name was needed since O. laciniata is the type of Raimannia. Since the complex connections between Renneria and Allochroa are now better under- stood, species of hybrid origin between them, all complex heterozygotes, have been designated as a distinct series here named Clelandia (11 species). The species here treated as series Clelandia have generally been treated in the litera- ture under the names O. odorata, O. stricta, O. indecora, and O. nana. The species of Renneria (in the sense of Fischer, 1962), with their subangular seeds and heavyset capsules, are confined to the high Andes. They probably are close to the ancestral form for the section, and may have subsequently given rise to the other South American groups. The species of series Allochroa occur mainly at lower elevations, although some species do extend into the moun- Ficures IA. Habitats of Oenothera sect. Oenothera taxa in South America.—1. Habitat of O. nana and O. punae, between Yaví and La Quiaca, 3,500 m, Jujuy, Argentina (Santarius). — 9. Habitat of O. indecora subsp. bonariensis, Itajaí, Santa Catarina, Brazil (Dietrich ).— 3. Habitat of O. catharinensis, Isle of Santa Catarina, Santa Catarina, Brazil (Dietrich ).—4. Habitat of O. stricta subsp. stricta and O. villaricae, Río San Pedro near Lago Rinihue, Val- ívia, Chile (Stubbe). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 429 430 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 tains. In the South American continent, various lines of the section have radiated into habitats ranging from coastal beaches to relatively high elevations. The common ancestor of these groups seems to have reached South America within the past several million years (Raven & Axelrod, 1974). It radiated ex- tensively in South America in relation to the many drastic changes in habitats that occurred in the Pleistocene and more recently (Simpson, 1975). The ar- rival of O. laciniata subsp. pubescens in South America was probably an event that occurred considerably later, perhaps only 100,000 years or so ago. CYTOGENETIC BASIS OF EVOLUTION Just as in North American species of subsect. Euoenothera, there occur among the South American members of sect. Oenothera species which are chromo- somally homozygous and form pairs of chromosomes or small rings at meiotic metaphase I, as well as others which are complex heterozygotes and form a ring of 14 or other large rings at meiotic metaphase I. Some of the plants which are essentially chromosomally homozygous also form small rings which segregate in an alternate fashion at meiotic metaphase I. All of the complex heterozygotes are highly self-pollinating as they are throughout the Onagraceae, with the possible exception of O. erythrosepala, whereas the chromosomal homozygotes range from self-pollinating to strongly outcrossing. All homozygotes of subsect. Munzia that have been investigated were genetically self-compatible. Most of the complex heterozygote species are true-breeding hybrids which combine genomes from two species which may be very distinct from one another. Their progenies are however uniform; they behave as true-breeding distinct species in nature. On account of the alternate disjunction of the ring of 14 chromosomes in such complex heterozygotes, two sets of 7 chromosomes behave as two dif- ferent chromosomes when segregating in meiotic anaphase I. In this way the pos- sibility of interchange between the two genomes and consequent segregation is minimized. Additional aspects of the genetic system, such as lethal factors, genes for self-sterility, and selective fertilization (Arnold, 1962; Schwemmle, 1968) pre- vent the reconstitution of the homozygotes at least in a viable form. Following self-fertilization, which is the rule in the complex heterozygotes, or outcrossing with another similar individual of the same population, the complex heterozygote is always reconstituted. Only in exceptional cases does one of the chromosomally homozygous parental types segregate in the progeny of a complex heterozygote, as discussed under O. acuticarpa on p. Cleland (1972: chap. 7) has given a 1 explanation for the origin of the rings of 14 chromosomes which occur in complex heterozygotes of Oenothera. The rings are formed because of reciprocal translocations between nonhomologous chromosome arms. Such reciprocal translocations, which probably occur at a low frequency in all groups of organisms but become a regular part of the adap- tive system in only a few, do not lead to an unequal distribution of the chromo- somes to the poles in Oenothera because of regular alternate disjunction of rings of chromosomes in this genus. All of the chromosomes are the same length, have 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 431 uRES 5-6. Habitats of Oenothera sect. Oenothera taxa in South 5 (continued). —5. Habitat of O. santarii, Cerro los Gigantes, 2,000 m, Sierra Gran Córdoba, Argentina ( Dietrich ).—6. Habitat of O. ravenii subsp. ravenii, $an Bernardino, D. Ypacaraí, Cordil- lera, Paraguay (Dietrich). arms of equal length, and appear euchromatic only at the end segments; this structure may facilitate alternate disjunction in the rings (Kurabayashi et al., 1962). Similar translocations between dissimilar nonhomologous chromosomes would lead to a high percentage of misdivision and therefore to a degree of sterility in the plants in which they occurred. In the evolution of the genus Oenothera the phenomenon of reciprocal translocation has played a decisive role. Following self-pollination or cross- ing of plants with small rings of chromosomes, it is possible to observe in the progeny not only chromosomal homozygotes which form 7 bivalents structurally identical to the original form of the species but also others which differ from the original form in the arrangement of their chromosome arms. Such plants constitute the starting point for the origin of new complex heterozygote species in the genus Oenothera. Through the hybridization of homozygous strains with others with a sufficiently different end arrangement, a ring of 14 or other stable configuration may arise, and then we can speak of permanent structural heterozygosity. In contrast to the North American species of subsect. Euoenothera, there are among the South American species many homozygous ones in addition to a rich 432 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 array of complex heterozygotes. Since all complex heterozygotes are by definition capable of being traced back to a homozygous original form, we have attempted in the course of our work to synthesize existing complex heterozygotes through hybridization between existing chromosomally homozygous species when pos- sible, and by crossing such complex heterozygotes back to their putative parents to illuminate the nature of the chromosomal complexes contained in them. t was possible to make a very comprehensive analysis of the complex heterozygotes in this group owing to the virtually unrestricted crossability of all species of this section. Fortunately, all of the genomes represented in the complex heterozygotes among the South American species are still represented in homozygous combinations among other species of the group. This should not be taken to infer that the phenotypes of these homozygous species are ab- solutely identical with those of the strains which originally gave rise to the com- plex heterozygotes, however, for many genomes have during the time they were combined in such complex structural heterozygotes become altered to a certain degree. though Oenothera does have an unusual genetic system, hybridization may play no greater a role in the origin of new species in this genus than it does in many groups of higher plants (e.g., Ehrendorfer, 1971). Only in the exact method of origin of the complex heterozygotes, which may perpetuate inter- mediate hybrid phenotypes, do they differ strikingly from methods of the origin of species through hybridization found in other groups of plants. DISTRIBUTION AND PHYLOGENETIC RELATIONSHIPS The general range of the South American species of this group and of the individual series is shown in Fig. 7. Especially notable is the extensive zone of overlap between Renneria and Allochroa as well as the frequency of chromo- somally homozygous species of both subsections in the vicinity of the Cordillera Oriental of Argentina. In addition, most species of the entirely hybrid series Clelandia occur in this area. The occurrence of so many species of both groups, and of the intermediates between them, in this particular region is probably related to the very close genetic relationship between Allochroa and Renneria, plants similar to the latter group having probably given rise to the original members of series Allochroa during the past. Rapid evolution of the species of the two groups in this region has no doubt been made possible in part by the rich diversity of habitats found in a relatively limited area along the east side of the Andes. Moreover, because of the impenetrable tropical forest and of the Gran Chacó, it is not surprising that the first migration out from the region of the Cordillera Occidental occurred towards the east and southeast into the open plains of Argentina. From here, secondary migration eventually resulted in the colonization of Paraguay, Uruguay, and southern Brazil. Ultimately members of this group migrated by way of the Andean passes to Chile. Species of series Renneria occur outside of the Andes only in the highest elevations of the Sierra Grande and the Sierra de Comechingones. Only their derivatives in which the characteristic phenotype of series Allochroa was evolved 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 4 JE | | Y T Ill Un JN " A Nt PL 1 1⁄4 2 7078 — — EHE ON. Hai Z 4 a 5 z sare L i 4 A 7 N S AS Ficure 7. Areas of the three series of subsect. Munzia: ser. Renneria, vertical lines; ser. Allochroa, horizontal lines; ser. Clelandia, diagonal lines. 434 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 were able to colonize the plains and eventually reach the seacoast. The species of series Renneria have remained strictly adapted to the high mountains. In Fig. 7 it can also be observed that the combination of genomes from series Renneria with those from series Allochroa, resulting in the origin of the en- tirely complex heterozygotes of series Clelandia resulted in plants which were able to greatly extend the area of the two parental series. On the one hand, series Renneria was able to penetrate to the southernmost point of the continent of South America at Punta Arenas only in the combination Oenothera magel- lanica, while on the other hand series Allochroa was able to penetrate to high elevations in the mountains of southern Peru only in the combination known as O. punae. Figure 8 is a plausible phylogenetic scheme for the evolution of series Allochroa from Renneria. Oenothera odorata and O. mendocinensis were de- rived from a plant which probably resembled the present day O. santarii, and they have remained confined mainly to the southern regions of South America. On the other hand, O. ravenii, O. longiflora, O. indecora, and O. affinis have extended their ranges mainly to the east. Oenothera ravenii and O. longiflora may have belonged to a group which developed around plants with a phenotype somewhat like that of O. longituba; O. indecora and O. affinis seem to be more closely related to a plant with the phenotype of O. scabra. Oenothera cathari- nensis seems to have been derived from O. ravenii as an obligate annual of local distribution. Oenothera verrucosa with O. coquimbensis and O. featherstonei seem clearly to have been derived earlier from plants similar to those of series Renneria, and independently from those species mentioned above of series Allochroa. Even though Munz (1935) put the taxa here regarded as series Allochroa and Raimannia into the same subgenus, it is difficult to establish any direct con- nection between them. On the other hand, there is a close and demonstrable relationship between the species of Allochroa and those of Renneria, treated by Munz (1935) as comprising separate subgenera. Their relationship is demon- strated by the fertility of almost any hybrid combination and even more strongly by the compatibility of their plastids. The hybrids between Renneria and O. indecora, O. mendocinensis, and O. odorata suggest that these species may be closest to Renneria, since they are capable of becoming fully green in both directions. In addition, albino or variegated plants have not been observed to occur among hybrid progenies involving these species with one another. In contrast, there are significant differences between the plastids of subsect. Raimannia and those of subsect. Munzia: hybrids between O. laciniata subsp. pubescens and species of Munzia were albino or very light green and set seed poorly (unpublished; Stubbe, pers. comm. ). There appears to be full plastome compatibility within the taxa of series Renneria, since all hybrids I have observed thus far were fully green. The estab- lishment of differentiated plastome types seems to have taken place within sub- sect. Munzia only after the evolution of series Allochroa from Renneria. Ac- cording to present information based on hybrids I have examined, it appears that O. affinis, O. longiflora, and O. ravenii each has its own distinctive plastid type. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA FicunE 8. Phylogenetic scheme of the evolution of Oenothera subsect. Munzia. 435 436 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 If this is the case, there would be at least four different plastid types within series Allochroa. Our investigations concerning the representation of these four plastid types among the complex heterozygotes are incomplete, and it is not possible to make a definitive statement about this matter at present. nlike the situation among the North American species of subsect. Euoeno- thera, distribution of the chromosomally homozygous forms in South America seems to have remained approximately as wide as that of their complex heterozygote derivatives, at least in series Renneria and Allochroa. Only in the cases of O. peruana, O. santarii, O. ravenii, and O. longiflora does it seem likely that the present area of distribution represents only a remnant of an earlier and much wider distribution. In addition, the ranges of O. verrucosa and O. featherstonei might be regarded as relictual. Whole series of species of Renneria and Allochroa seem to have acquired the ability to greatly extend their ranges of distribution, in a sense, by form- ing new complex structural heterozygous combinations with other species. From this it can be assumed that the ability of these complex heterozygotes to colonize new areas is greater than that of the corresponding homozygotes. This seems to be especially clear with respect to Chile, where only complex heterozygous species of subsects. Allochroa and Clelandia occur, except, of course, for the occurrence of O. coquimbensis in the deserts of the far north. In the course of their migrations the complex heterozygotes seem often to have migrated by a circuitous path, as can be illustrated especially clearly by the history of the O. odorata complex. The odorata-complex first of all participated in the origin of O. stricta, the other parent being O. ravenii. Oenothera stricta is now very widespread and abundant in Chile. Figures 241-244 show other examples of the spread of different complexes. ANALYSIS OF COMPLEXES The analysis of complex heterozygotes in Düsseldorf was made possible by the comparison of various artificially produced complex heterozygotes with naturally occurring ones. In addition, naturally occurring complex heterozygotes were hybridized with their putative chromosomally homozygous parents in both directions, and the hybrids evaluated morphologically. The association of chromosomes was of no value in most cases in assessing the relative parentage of the different forms. Haustein (1952) pointed out over 20 years ago that chromosomal pairing was of almost no value in determining phylogeny in this group, as it was always complete and variable within many species. If, for example, a hybrid between a randomly chosen line of Oenothera versicolor and one of O. scabra formed 7 bivalents, one could certainly con- clude that these two strains are related to one another, since the pairing of their chromosomes is undisturbed, and fertility, whether measured by pollen or seeds, is complete. About the degree of relationship between them, however, relative to other species of the group, this test says nothing. In many instances, the evolution of species within series Renneria especially seems to have taken place solely by genetic changes, not involving any rearrangement of chromo- 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 437 some structure. As another example, I might mention that “longiflora Erlangen,’ a strain of O. affinis (series Allochroa), gave a hybrid with a ring of 4 and 5 bivalents when crossed with a strain of O. scabra (Santarius 2003; series Ren- neria) collected in Bolivia. These structural homologies cannot be taken as a strong indication of a particular relationship between O. affinis and O. scabra, however, since within O. affinis itself, hybrids have been observed to form configurations at meiotic metaphase I ranging from 7 bivalents to a ring of 14. Other indications that chromosomal configurations in hybrids between various South American species are not a useful index to relationship is provided by the following observations. Even in individual populations of O. affinis and O. odorata—populations which appear morphologically homogeneous—there may be found individuals forming 7 bivalents, others which form a ring of 14 at meiotic metaphase I, and still others with smaller rings. In short, chromosomal evolution does not seem to have proceeded among this group of species at the same rate as morphological evolution, and the two do not appear to be highly correlated. Variation of this sort has been found in limited areas; for example, Santarius 1869 represents a sample that in the experimental garden was mor- phologically indistinguishable from Santarius 1850, and both were referable to O. affinis. When hybridized, these two populations yielded progeny in which the plants formed a ring of 14 at meiotic metaphase I. On the other hand, when plants from Santarius 1869 were hybridized with a morphologically very dis- tinctive strain with broad leaves from 1,600 km away—Santarius 193—they formed a ring of 4 and 5 bivalents. When they were hybridized with Santarius 1711, another strain of O. affinis from about 350 km away, they formed a ring of 6 and 4 bivalents. Oenothera affinis is evidently in a stage of evolution in which the chromosomal homozygotes are more widespread and much better represented than the hetero- zygotes. The sorts of intermediate chromosomal configurations that occur fre- quently indicate, however, that a wide variety of different end arrangements occurs in the species. In cultivation, the complex structural heterozygotes breed true and do not “throw off” homozygous derivatives. This pattern is very different from that worked out by Cleland (summary in Cleland, 1972) for the North American subsect. Euoenothera. On the basis of his studies of this group, Cleland worked out a theory whereby the accumula- tion of reciprocal translocations within a population, accompanied by geograph- ical isolation, would lead eventually to the origin of populations which were essentially chromosomally homozygous but characterized by different end arrangements. If these strains eventually spread and came into contact, com- plex heterozygotes might originate in a single step by hybridization in the zone of contact. These complex heterozygotes, fixed from their origin, would breed true and would eventually replace the chromosomal homozygotes with which they occurred. A situation similar to that just described for O. affinis can be accepted for O. odorata. With respect to its chromosomal configurations, O. odorata is ex- tremely variable. Plants with 7 bivalents and all configurations up to and in- 438 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 cluding complex heterozygotes with a ring of 14 occur. From this, it can be concluded not only that a variety of differentiated chromosomally homozygous types occur within the species, but also that the species is still in a state of active evolution chromosomally. It is not clear whether the complex heterozygotes within this species have formed through hybridization between chromosomal homozygotes completely differentiated with respect to their end arrangements, or have accumulated following the formation of rings of chromosomes of in- termediate size by further spontaneous reciprocal translocation. In some pop- ulations, the balance seems to have shifted definitively toward a preponderance or exclusive representation of chromosomal structural heterozygotes, while in others, chromosomal structural instability seems to be the rule. In the vicinity of Comodoro Rivadavia, Prov. Chubut, the essentially stable configuration of a ring of 12 and 1 bivalent seems to have become predominant. Results such as those discussed for O. affinis and O. odorata, lead to a view of the origin of complex heterozygosity quite different from that presented by Cleland and others for subsect. Euoenothera. In these South American species, as in some others, chromosomal structural heterozygotes occur together with homozygotes and plants with intermediate configurations in the same popula- tions. There is evidently no direct correlation between the formation of strains or populations that are well differentiated either genetically or phenotypically and the origin of complex heterozygosity, since in these cases the only major dif- ference between the genomes represented in the heterozygotes seems to be that concerning the actual end arrangements represented. It seems unimportant whether the pattern of end arrangements responsible for the ring of 14 arose gradually or in a single step. Oenothera affinis, like all other South American species of the section, is self-compatible and often self-pollinating, the anthers shedding pollen directly on the stigma at anthesis. Therefore, regardless of how a complex heterozygote originates, it would tend to persist in the population alongside plants with other chromosomal configurations owing to regular self-pollination. This species exhibits several stages of complex heterozygosity. The first, in which homozygotes still segregate from the heterozygotes, is not known in O. affinis, although it does occur in O. acuticarpa. In the second stage, the com- plex structural heterozygotes are plants with large flowers and a long floral tube, indistinguishable from chromosomal homozygotes of the same species. The population of O. affinis in Chile is probably nearly or quite dominated by complex heterozygotes. However, only one bit of actual cytological information is available (Stubbe in 1961). Examination of the pollen of plants from four populations from the provinces of Atacama, Coquimbo, and Valparaiso (John- ston 5860; Behn 8468, 22798; and Eyerdam 10040) revealed from 20-35% empty grains, however. A fifth population, from Aconcagua (Behn 22802), had essen- tially no empty grains, which makes it virtually certain that it was chromosom- ally homozygous. The origin of complex heterozygosity in the Chilean popula- tions seems to have been very recent, and it may be that the original plants that became established on the west side of the Andes were already heterozygotes. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 439 The third phase in the evolution of complex heterozygosity occurs when all plants are small flowered and self-pollinating, although retaining the long floral tube characteristic of O. affinis. Further reduction of flower size would lead eventually to plants with a short floral tube also, as has occurred in the evolu- tion of O. mollissima from O. affinis. The morphological changes that have occurred in O. mollissima have been so extensive that it is best regarded as an independent species at the present time (Hecht, 1950; Hecht & Tandon, 1953). Even though these species often occur in mixed populations, the regular self- pollination of O. mollissima seems to constitute an effective and sufficient bar- rier to interspecific hybridization in nature; hybrids occur only occasionally. The origin of complex heterozygosity in O. elongata (series Clelandia), in contrast to the situation discussed in O. affinis and O. odorata, involved N ization between two distinct taxa. Other clear-cut examples of this sort of evo- lutionary process are afforded by the origin of O. magellanica, O. siambonensis, O. stricta, and O. villaricae. The strain of O. elongata from the province of Catamarca (Diers in 1959) has had a remarkable history, quite apart from the fact that the locality is far from the main area of distribution of the species. In 1960, an individual plant appeared in the progeny of a seed sample from O. longituba at the same locality, gathered in the wild. In all probability, this plant was a spontaneous hybrid between O. longituba and O. affinis, both of which commonly grow together in Catamarca. On self-pollination, however, it bred true, and it has now been maintained in cultivation for 13 years as a com- plex heterozygote, without showing any tendency to segregate the characteristics of its presumed parents. The original plant gave evidence of plastid incom- patibility and probably would not have been vigorous enough to survive in the wild; by a fortunate chance, it was grown in the experimental garden to give clear evidence of the mode in which a particular complex heterozygote could have been formed. Evidently the one-step, efficient formation of this chromosomal arrangement deterred the segregation of the parental species from the start. Spontaneous hybrids with O. affinis also appeared in the progenies of two other collections of O. longituba, Santarius 1736 and 1737. These had inter- mediate chromosomal configurations, however, of a ring of 6, a ring of 4, and 2 pairs; 2 rings of 6 and 1 pair; and a ring of 12 and 1 pair, and probably would not have been stable as was the plant discussed above and assigned to O. elongata. Two principles seem to have been important among the evolution of different South American species of this group. In some species extensive genetic changes have taken place without structural modifications of the chromosomal end ar- rangements. Thus, the strains of O. peruana, O. versicolor, and O. scabra that have been grown in the experimental garden have been chromosomally structurally identical. Other species of series Renneria have not deviated far from this chromosomal structural arrangement either, so that a ring of 4 and 5 bivalents was formed both in hybrids between O. santarii and O. versicolor and also in hybrids between O. pedunculifolia and O. versicolor. In many other species structural changes in the form of reciprocal trans- locations have played a definitive role in the differentiation of species. These 440 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 translocations have not led to a reduction in fertility because of the regular alternate disjunction of the chromosomes in the rings. As a result, a whole ar- ray of chromosomally distinctive lines has arisen from such events. Familiar isolating mechanisms of a geographical, ecological, or physiological nature have in many instances allowed complete speciation following or accompanied by such chromosomal differentiation. In the absence of barriers to hybridiza- tion between chromosomally differentiated lines within the group, many such lines have eventually hybridized to produce new complex heterozygous com- binations. Indeed, identical combinations have been found, evidently formed more than once at different times in different places. Examples of this are pro- vided by O. stricta and O. picensis, and these are discussed further under the treatments of the respective species. As in the chromosomal homozygotes, genetic changes that have occurred in the complex heterozygotes have led eventually to a still further differentiation. The tendency towards the evolution of small-flowered plants in which self- pollination is more and more automatic, regarded in this paper as the ultimate phase in the development of complex heterozygosity, is unmistakable. Despite the extensive suppression of crossing-over in Oenothera, the occasional ex- change of genes between complexes seems to have had an effect on the course of evolution. An example of this, O. elongata, with complexes derived from O. longituba and O. affinis, has been mentioned. Hybrids between this complex heterozygote and O. affinis have shorter capsules than O. affinis and this char- acteristic of O. longituba seems to have been introduced by gene exchange in the complex heterozygote. In contrast, O. picensis (Santarius 1549) is clearly made up of more or less unaltered O. affinis and unaltered O. odorata. Also, O. villaricae, which is known as “berteriana Erlangen" in the genetic literature and belongs to series Clelandia can be shown to be made up of complexes de- rived from O. santarii and O. ravenii. Among the additional factors of phylogenetic importance among the South American species may be mentioned complex heterozygosity involving more than two species (see species numbers 12, 16c, and 28). A very curious phe- nomenon involving O. nana and O. punae, involving a situation in which freely recombined genomes at a single locality seem to be unequally shared by both species, is discussed on p. 611. An important result of this study is the conclusion that the origin of complex heterozygosity does not depend upon the prior existence of genetically well- differentiated genomes. It can be shown, especially in O. affinis, that chromo- somally very well-differentiated forms can occur within a single population, be preserved by self-pollination, and eventually give rise to complex heterozy- gotes which are morphologically essentially identical to the chromosomal homo- zygotes at that place. (See also Drillisch, 1975: 60-71.) Such complex heterozy- gotes, since they are from the beginning self-pollinated, may immediately be more or less isolated from other elements of the population and thus in effect give rise to a new and distinctive strain. Thus rings of 14 can arise in a variety of ways in populations of Oenothera, either through the hybridization of chro- 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 441 mosomally and genetically very well-differentiated strains, or through the grad- ual accumulation of differences, even within populations. In contrast to the species of the North American subsect. Euoenothera, chromosomal homozygotes are still very well represented among the South American species, even though complex heterozygotes are likewise frequent. These chromosomal homozygotes still have ranges as extensive as those of their complex heterozygous derivatives, and often occur together with them in the same areas (Table 1). The analysis of the South American complex heterozy- gotes is facilitated by the existence of all of the original participating homozygotes at the present time. The various mechanisms operative in the differentiation of species all can still be observed directly at the present day. n very actively evolving groups, such as the species of Oenothera in South America which are treated in this paper, it is often not possible to delimit sharply defined natural units. Almost all complexes of series Renneria and Allochroa are interconnected with one another, and they are in a very real sense evolving as an interconnected whole. Connections between species, taken step by step, unify the most distantly related and distictive elements in the group into an inter- twined complex of great intricacy but presumably also great evolutionary vigor in responding to the demands of the environment. The two most im- portant elements in the evolution of the group, however, appear to be com- plex heterozygosity and hybridization. This seems to be particularly true among the South American species of Oenothera because of the complete lack of sterility barriers, and because of the chromosomal system which leads easily to the formation of true-breeding hybrids which recombine the characteristics of the parents in various adaptative ways. The frequent instances of sympatric occurrence of the various taxa, with consequent opportunities for hybridization, is documented in Table 1. In fertile, pair-forming hybrids such as are encountered in many groups of plants, adaptive phenotypes may soon be broken up by recombination and back- crossing, but in stable hybrids, such as self-pollinating complex heterozygotes, it may be preserved. Once a successful complex heterozygote has originated, it may spread in areas beyond those occupied by its homozygous parents, owing to its different adaptive mode, and eventually the parental homozygotes may come to exist only in relatively limited areas, as in the North American subsect. Euoenothera. SysTEMATIC TREATMENT Before commencing the formal taxonomic treatment of the South American species of this group, I would like to make a few comments about the descriptions and the way I have employed terminology in them. For the descriptions of in- dividual pubescence types the terms strigillose, villous and glandular-pubescent have been used. All hairs are fundamentally single-celled. "Strigillose" is used to designate short, appressed hairs about 0.1-0.2 mm long; “villous,” hairs which stand erect or rise obliquely and are 1-5 mm long; and “glandular-pubescent,” hairs 0.1-0.3 mm long which are terminated by a gland. Shorter and appressed 442 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 l. Sy EN occurrence of taxa of native South American taxa of Oenothera sect. TN aes subsect. Mun ME = ° clolS} |s BEER HE 2 slal EEE MEBBHHEMNEERBSESRERSBBSBEBEBMBENMEHBHSEHHEOSHHREEBHBREHHEBHENE HEHHBBBSSBHHRHHBBBBSEREBEERSHEHHHBEHSESHHBHEBHEEHSEBHBBE 9 2 4E = z E S15 = dE dB BBBBBEEBBEBBH EHE BE EE BH E B EHE EE REN ruana ° ° —— e| jejejeje ele ° 2 Tasiocarpa | e 888 ° ° ° santarii ° ° le ° Tongtuba — jeje goc è g e| je] fe tafien ta 888 ele e e ° eje ° ° pedunculif eje| jeje U ° ° ° ejeje scabra Jeje] jejeje D g e[e| je] Jo] je rubida le | . tarijensis ° ° ejeje ° ° ° Tecurva ° ° ele ° . ° sandana ° e| Jojo] je dg ete nana ele] jejejeje| jejeje Ü . "LL mendocinen è e[e| Jè 28888 ele e “odorata . ° ° e| jejejeje| je ° ° Taveni rav — e| [e[e[e[e e| [e| Je Taveni arg g e| jejejeje ele! jejeje] jejeje raveni chi ° ° longif gra ejeje| | e| je longif lon LJ lele le ejeje ° catharinen Deco ind ele ojojo e| je| je| je 2 — indeco bon u eje eje ° eje [| je| | je) eee ° ° affinis ej[ejejejejojo eje e| jeje eje eje e ° ° eje| jejeje ° e| je e| je eje ejeje “mollissima | Je] jejeje ejejeje| je rivadaviae eje ° ° strict str l ° | [e g LI e| je eje strict alt . II — 1 + J eje strict arg eje] je jeje ele eje bahia-blan ele] je LI ° ° 2 picens pic eje ] Je ° ej je ° picens cor ° ° gog . U ° ° “picens bon - eje] [o ° ele ele “monteviden eje elelele e[ [e pseudolong ° ee a parodi par e| jeje U ejejejeje| | eje ejejo eje ° e parodi str_ e| je ele ° ° parodi bra eje eje e. eje e ° verrucosa ° ° | Coquimbens > ° Ogi ° ° arequipens 1 gg g grisea e. e| | ° ° feathersto | ° g ° nocturna ° ° ° tre magel anic ° ° eje ° eje gogg . eje villaricae ° eje e elongata ° e| lelelelo pseudoelon . ° ° ° cordobensi e ° ° ° . siambonens ejejejojo ° ° eje brevipetal ° ° acuticarpa eje ° ° . tucumanens e eje ° ° ° punas e[e| |ejele]e| jejejeje ° "RAP Tacit pub O e| jeje villous pubescence is likewise encountered among the South American species of Oenothera. All three pubescence types may be present on a single plant. Description of the form of the vegetative leaves and petals, as well as that of the outline of the buds, capsules, and seeds follows the terminology given by Stearn (1956). Measurements for the buds are given for those just about to open. The length is given exclusive of the floral tube. Measurements of the sepals are given at the same stage. Length of the ovary is given at the stage when the flowers have just opened. The asterisks on the chromosomal configurations mentioned at the end of the description correspond with those given in the lists of plants cultivated in the experimental garden at Düsseldorf. The geographical distributions are in general given in broad terms, since many of the ranges, especially those involving the species occurring in the high 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 443 mountains, are insufficiently known. Further sampling will be necessary before they can be specified more precisely. Finally, it is important to add a word about the duration of the plants. Often in the literature and on the labels of specimens, plants of this group are said to be “perennial” In most instances I have not been certain how to treat these observations, since in our experience all species appear to be either biennial (especially the rosette-forming ones) or annual. Probable exceptions, and spe- cies that may truly be perennial, are O. nana and O. punae. I have not been able in any case to find on a herbarium specimen any plant parts indicating that the individuals had flowered more than once, or structures such as over- wintering buds which might have indicated a true perennial habit. Structures which might have seemed to have been formed during an earlier flowering cycle could in all cases be attributed to the activities of sheep or other grazing animals. There seems to be no indication that any of these plants are generally perennial, although when they are cropped off by an animal or cut back in a stage of active growth, new shoots will often form freely from the base and a second flowering will take place within the same season. In Diisseldorf we often employed this device in order to synchronize the blooming of two different species which might not otherwise have flowered at the same time. OENOTHERA sect. OENOTHERA Three subsections are represented in South America, and may be separated by the following key: 1. Seeds prismatic; infrequent DUUM plants C. Subsect. Euoenothera (p. 615) l'. Seeds not prismatic; mostl Buds nodding or floral vitae of the oldest buds curv 1 upwar ra. ubsect. Raimannia (p. 612) 2’. Buds erect, the floral tubes not curved upward... A. Subsect. Munzia (p. 443) A. Subsection MUNZIA Oenothera sect. Oenothera subsect. Munzia Dietrich, subsect. nov. Plantae annuae vel biennes, erectae vel prostratae, rosulares vel erosulares; gemmae non nutantes, tubus floralis non sursum curvatus. Habitat in America Meridionalis. Annual or biennial plants, erect or prostrate, forming a rosette or growing directly from seed without one, or remaining permanently and flowering as a rosette. Plants unbranched or with a branched main stem and decumbent, arch- ing, or straight side branches arising from the rosette, a few cm to 2 m tall. Rosette 5 cm (O. nana) to 70 cm (O. longituba) in diameter. Main stem 0.3-2 cm thick. Plants (1) very densely to sparsely strigillose and thickly to sparsely appressed to erect long-villous; (2) very densely to sparsely strigillose, densely to sparsely long-villous, and very sparsely to sparsely glandular-pubescent; (3) very densely to very sparsely appressed or erect long-villous, densely to very sparsely glandular-pubescent; or (4) densely short-villous and glandular-pubes- cent. Rosette leaves linear, elliptic, or oblanceolate, long or short acute, gradu- ally narrowed to the petiole or sessile, attenuate to acute at the base, 2.5-35 cm long, 0.2-6 cm wide; cauline leaves linear, elliptic, lanceolate, or oblanceo- 444 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 late, acute, sessile, acute to truncate at the base, or distinctly short-petiolate and narrowly cuneate at the base, 2.5-25 cm long, 0.1-4 cm wide; bracts linear, elliptic or lanceolate to ovate, acute to almost obtuse, sessile and acute to subcordate at the base or distinctly short-petiolate and attenuate at the base, l- 12 cm long, 0.1-3 cm wide, longer or shorter than the capsule they subtend or + the same length; leaves regularly or irregularly to sparsely toothed, the teeth obtuse or acute, plane or undulate, the surface sometimes flecked wih black (O. nana). Inflorescence simple or branched, dense or rather sparse. Floral tube 0.5-12 cm long, sometimes flecked with red or reddish brown. Buds oblong, elliptic to rotund or lanceolate to ovate in outline, 0.4-3.5 cm long, 0.15-1 cm thick, often red striped at the junction of the sepals or also along the midribs of the sepals. Sepals green or yellow to greenish yellow, often flushed with red, sometimes flecked with dark red or reddish brown; sepal teeth 1-4 mm long, erect or divergent. Petals broadly elliptic or obovate to broadly obovate, rotund or retuse, 0.54.5 cm long, yellow to straw colored, sometimes flushed with red near the base and along the nerves, or completely red. Anthers 0.2-1.5 cm long, yellow, sometimes reddish. Filaments 0.2-2.5 cm long, yellow, sometimes red- dish. Style 1-14 cm long, surrounded by the anthers in bud or exceeding them in length. Stigma lobes 4, 1-10 mm long. Ovary 0.8-1.5 cm long, cylindrical. Capsule ovate or oblong in outline, narrowed toward the apex, occasionally swol- len below the apex (O. affinis), fused with the subtending bract at the base or free, occasionally clearly stalked (O. verrucosa), erect and appressed to the stem or spreading sharply or even at a right angle to it, 1-6 cm long, 2-10 mm thick; valves of the capsule separating from the placenta when ripe, spreading or curving inward or outward. Seeds 0.8-2 mm long, 0.5-1 mm thick, = bluntly angled or elliptic to rotund in outline, light brown to dark brown or black, sometimes dark flecked. Self-compatible and outcrossed or self-pollinated. Chro- mosomally homozygous or permanently heterozygous and isogamous or semi- heterogamous. Gametic chromosome number, n — 7 (7 bivalents, ring of 14, or intermediate configurations at meiotic metaphase I). Type species: Oenothera odorata Jacq. Distribution (Fig. 7): All species occur in more or less open plant com- munities from sea level to almost 5,000 m elevation in the Andes (O. nana). mong the sorts of places they grow are seacoasts, dunes and other sandy places, gravelly fields, dry watercourses; fields, meadows, and pastures; open shrublands and woods; grassy and shrub-steppes, Andean puna; banks of streams; edges of roads and disturbed places. The subsections occur nearly throughout the southern half of South America, and northward along the Andes to Colombia; some of its species are widely cultivated and naturalized on other continents. This subsection is dedicated to P. A. Munz (1892-1974), lifelong student of Onagraceae. KEY ro THE SERIES or SuBsECT. Munzia 1. Capsule urn-shaped (Fig. 9), gradually narrowed toward the tip, erect or standing out at a right angle from the stem, 4-9 mm thick at the base and with the subtending 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 445 FicunEs 9-12. Capsules representative of the series of Oenothera subsect. Munzia.—9. O scabra subsp. scabra (Renneria).—10. O. rivadaviae ( Allochroa).—11. O. acuticarpa (Cle- landia).—12. O. verrucosa ( Allochroa). leaf distinctly fused to it; valves of the capsule spreading after seeds are shed MESURER INN Series oid Ap. 450) . Capsule + cylindrical (Figs. 10-12), at most tapering toward the a Or standing out from the stem, 1.5-4 mm thick, usually not or slightly led 21 the subtending bract; valves of the capsule spreading or curved inward or outward after seeds are shed. Capsule cylindrical (Fig. 10), rarely enlarged in the upper third or somewhat petiolate, not fused with the subtending brate Series nw = 489) '. Capsule generally gradually narrower i. vard from a broad base (Fig. dently fused with the subtending bract for a short distance .. Series E odd e 585 , — to KEY ro THE SPECIES OF Oenothera SECT. Oenothera IN SOUTH AMERICA l. Seeds prismatic Subsect. ne 2. Floral tube 2-5 cm als 3.5-5 cm im MR ORC TERRENCE EORR EB 48 E 5 š als 1-2.5 cm long ————— I O. bie p. 618) 2’. Floral tube 1-1.8 cm long |... . 49. O. villosa i. strigosa l'. Seeds not prismatic. 4. Buds nodding or = pn tube of the oldest buds curved upward Subsect. (Raimannia). 5. Petals 0.5-1.5 c 6. md often ‘flaked with red; apices of the sepals 0.5-1 mm long SUYO A A 6a. O. laciniata subsp. pubescens Sep als never r flecked; a apices of the sepals 1-2 mm lon TE Sie tense asec s 2 EEE AE 46b. O. laciniata ws lacin "C Petals 2.5-5 em long S S l 47. O. a a ds not nodding; floral tubes not curved u upward Subsect. ( Munzia igh mountain plants of very condensed habit, either flowering in “the rosette or forming short, prostrate side branches; capsules 1-2 cm lon 8. 1 forming a rosette only, rarely with short side bes capsules ing out at right angles from the stem _ . O. nana Pow with prostrate side branches 5-20 cm sei capsule + appressed to the stem ̃ĩ⅛ů —— 45. O. punae 7’. Plants neither rosette-forming nor prostrate, or taller than 15 cm, erect or prostrate, but if prostrate, then the capsules more than 2 cm long. 446 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 9. Capsule urn-shaped, gradually narrowed toward the tip, erect, 4-9 m thick at the base and with the subtending leaf fused to it; valves of the esit spreading after seeds are shed; exclusively mountain plants (series Renneria in large part 10. Floral tube 6-10 cm u Toig: flowers exceeding the apex of the stem in ength. 11. Plants forming a rosette, with a crowded inflorescence; pubescence strigillose and erect villous; petals 2.5-4.5 cm long -.... O. longituba 11’. Plants not forming a rosette, with a more "Pe der and lax inflores- cence; pubescence strigillose and appressed villous; petals 2-3 cm lon 11. O. recurva 10’ iv tube Seed cm long; flowers mostly not exceeding the apex the stem in len 19. Floral tube 2.5- 55 cm long Plants with appressed “and erect pubescence, sometimes also glandular-pubescen 4. Buds narrowly lanceolate to oblong in outline; petals 2-3.5 cm long; seeds mostly flecked with m spots, 0.8-1.3 mm 14'. Buds lanceolate in outline; petals 15-25 cm ~ anes ab most always immaculate; plants never glandular pubescent 12. O. sandiana 13'. Pubescence all appressed. 15. Upper bracts EUR red margins; petals often reddish near the base and along the veins; seeds dark brown to bow black, 1.3-1.5 mm i. . 0. O. tarijensis 15’. Bracts without red margins; petals entirely yellow; 1 ET brown to dark brown, flecked with reddish brown, 0.8-1. mm long. 16. Leaves regularly and ird serrate; pass evidently petiolate; seeds 0.8-1 mm long 5 16˙. Leaves irregularly and usd serrate; Ds Rin n s 1-1.2 mm long 6. O. tafiensis 12’. Floral tube 1.2-2.5 cm long. l7. Plants with both appressed and erect pubescence. 18. Plants 1-3.5 dm tall; floral tube 1.5-2 cm long; pu 1.5-2 cm long, reddish at the base and along the veins . O. lasiocarpa je. ts 5-12 dm tall; floral tube 2-2.5 cm long; sel 2-3 ng, entirely yellow O. santarii m lon Ir. Plants with appressed pubescence only. 19. Bracts reddish along the Eug ie peta e red or with a red ipee at the base and reddish along the veins _..... 9. O. versicolor 19'. Bracts not reddish; petals entirely ies 0. Plants a a rosette; petals 1. 5-2.5 cm long; capsule mm thic O. peruana 20’. ii E forming. a rosette; petals 0.8-1.2 cm long; c cap- 4-5 mm thic O. Sum 9'. Capsule + cylindrical, at most tapering toward the apex, erect or standing out from the stem, 1.5-4 mm thick, rarely somewhat thicker, usually only indistinctly fused with the subtending leaf if at all; valves of the capsule spreading or curved inwardly or outwardly when ripe after seeds are shed. 21. Floral tube 7-11(-13) cm long. 22. Plants forming a rosette; pubescence thick and shaggy; 5 shorter than the capsule they subtend 17. O. longiflora 22’. Plants not forming a rosette; pubescence soft; bracts longer ina the capsule they subten 23. Bracts cultrate to narrowly lanceolate; capsule 2.5-4 cm long, 3-4 mm thick, somewhat enlarged in the upper third; seeds 1.5- 2 mm long 20. O. affinis 23'. Bracts lanceolate to narrowly ovate; capsule 2-3 cm long, 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 447 ana eS coe te LEE EER PRECIO TES EN . O. elongata 21’. Floral tube at most 6.5 cm long. 4. Capsule narrowed at the base and at the apex, apparently stipitate; valves of the capsule spreading when seeds are shed; annual plants, mostly less than 3 dm tall. Plants tnéstly ia m leaves 1 577 and bluntly serrate; floral tube 0.6-1.1 cm long; seeds 1.5-1.7 mm long, m browr n to almost Tk o ee LE O. verrucosa 25'. Plants branched; leaves sinuate-toothed; floral tube 1- 3 cm long; seeds 1-1.3 mm long, brown 9L. 0. es 24’, ae not narrowed at ilie base; valves curving when seeds s 26. Leaves with long, narrow teeth, these often secondarily toothed, or leaves remotely and bluntly serrate; valves of the capsule curving inward when seeds are shed; seeds RC elliptic in outline, 1.2-1.6 mm long, 0.4-0.5 mm thick O. coquimbensis 26' . Leaves without long, narrow teeth; valves of the que sule curving outward or spreading when seeds are shed; seeds elliptic to broadly elliptic in outlin 7. Plants with exclusiv ely appressed pubescence. Petals 2.5-4.5 cm long 33. O. featherstonei 28’. Petals less than 2.5 cm long. 29. Bracts linea 3 14. O. mendocinensis 777 4. O. bahia-blancae 30’. Capsule less than 3 cm long, 2— T. mm thick. 3l. Capsule shorter than the subtending br 6 28b. O. parodiana 5 strigulosa 31’. Capsule longer Han the subtending bra 3 Bracts lanceolate to o vate, truncate x su cordate at the base; leaves with Pioni teeth: seeds ain. in butline, 1.3-1.5 mm long 0.5-0.7 à Riek —- 32. O. „grisea 32’. Bracts 3 elliptic to p attea- uate at the )ase; leaves ostly sinuatr - Pese seeds broadly elliptic in outlire, 1.3-1.8 mm long, 0.8-0.9 mm thick _ 8 - 34. O. nocturna 27’. Plants with some erect ‘pubescence, _ Bracts linear o 14. O. mendocinensis Petals 2.5-5 cm long. 35. Bracts narrowly oblong to lanceolate, rounded at the base; plants not rn a rosette laal, MS 18. O. catharinensis 35’. Pade lanceolate. - to uncle uate: truncate to subcordate at the base; plants iini: a rosette. 36. Bracts longer than the capsule they subtend rep: 15. O. odorata 36'. Bracts shorter than the capsule they subtend, or at most subequal to it 37. Bracts usually only half as long as the cap- sule they subtend, or even shorter; floral tube (3-)3.5-5.5 cm long; capsule 2.5-3.5 cm long; Sud 1:315 mm len ER DON NOUS . 16a. O. ravenii subsp. ravenii 37'. Bracts more than half as long as the cap- sule they subtend; floral tube 2.5-3.5 cm ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 i capsule 3.5-5 cm long; seeds 1 og 93b. O. stricta subsp. 8 34’. Petals at ed 2.5 cm long. 38. Capsule 1-1.5 cm Ds „ 38˙. Capsule longer. 39. Capsule 1.5-2.5 mm thick. 40. Plants exclusively short-villous and dular-pubesce M appearing glabrous S viewed witho 190 42. O. brevipetala Non » O. 3 subsp. bonariensis 40’. age with erect, long villous pubescence 41. Tannie 3.5-6 cm long 22. O. rivadaviae 4l’. Capsule at most 3 cm lon ng. 42. Bracts only half as long as the cap sule they subtend or shorter, pee to subcordate at the base 8a. O. parodiana subsp. parodiana 42%. Bracts subequal to or longer than the capsule they subten 43. Plants strigulose, villous, and glan- dular-pubes 5 ect thea he funr on — O. E | Plants only villous and glandu- ar-pubescent; bracts subequal to r shorter than the capsule they subtend. 44. Plants forming a rosette, + o = — with dark red; capsule 1.5-2 mm thick; seeds 1-1.3 mm Ong c LLL 19a indecora subsp. indecora 44'. Plants not forming a rosette, densely villous; sepals not Yes with p 5 2— = m thick; ink. 26. 0. plait NR 39'. Capsule 3-4 mm thick 45. Bracts longer than the capsule they subtend. Plants only villous and glandular-pubes- cent; capsule standing out obliquely from the stem. ‘ d : Bracts rounded to truncate at the stem erect and somewhat imbri- cat 21. O. mollissima . Bracts truncate to subcordate at the base, spreading . 25. O. picensis 47’. Plants not very densely long-villous 25. O. picensis — oc 46'. Plants strigillose and villous; in part also glandular-pubescent; capsule + erect. Plants densely long-villous; 0.8-1.3 cm long .. 39. O. pseudoelongata 49'. Plants with various 5 e; pet- als at least 1.5 em long. 50. Plants pace a rosette; floral 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 50' they Ct * 449 de 1.3-2.5(-3) cm long; seeds immaculate. l. Bracts evidently longer than the capsule they subtend without JE margins; capsule 9 ong; seeds 1.4-2 C dec. 5- O. . Bracts Abii longer than Ot — 1.1-1.5 mm long 36. O. villaricae Plants not forming a rosette; floral tube 3-6 mm long; seeds flecked with reddish brown. 52. Bracts only a little longer than the capsule they subtend or s horter, mostly with red mar- Ea s 41. O. siambonensis i Bracts evidently longer than the capsule th zu subtend, without red margin 43. 0. acuticarpa 45’. Bracts subequal —- Or shorter than the cap- sule ubtend. 53. Capeuia suber 54. Pla ect. nts 5 5 5 a rosette; seeds im- maculate. 55. Floral tube 2-3 cm long 55“. Floral tube 44.5 cm long 33233 . O. villaricae EE. INS 37. O. hechtii 547. Plants not forming a rosette; seeds flecked with reddish 56. brown. Floral tube 1.5-2.5 cm LE cap- sules with the 4 valves evidently free and crenate at the apex EN. 40. O. cordobensis Floral tube 3-5.5 cm long; cap- sule without evidently free valves NM ` . siambonensis 53’. Capsule standing out from the stem 57. Petals at most 1.5 c an ong. ts not Sarees a rosette; floral tube bs cm long picensis subsp. cordobensis oral 9! O. 58’, 8 forming a rosette; tube at most 3 em EL ng. 59. Capsule 3-4.5 cm long — 24. O. ie P blancae 9’, Capst ule 2.5-3 cm long. 60. Floral tube 1.8-3 cm long; yuds 10-17 mm longi pet- d 1.2-2 cm long _.... `. - 16b. O. ravenii subsp. inae A 577 5 tube 1-2 cm long; D Q TON a. O. parodiana subsp. parodiana 450 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 57’. Petals more than 1.5 cm long. 61. Floral tube more than 4 cm long. 62. Floral tube usually more than 4. ) cm long; a. nar- rowly oblong to oblo in outline 27. O. pseudoloneiflora u . Floral tube at most 4.5 bes buds lanceolate in Ea ine 63. 63’. Upper bracts with arched incurved apices; plants with long, "bristly hairs, especially below ------------- 28c. O. parodiana subsp. brasiliensis Upper bracts plane; plants strigillose, especially be- low, never with long, stricta 8 stricta . Floral tube less than 4c E Bracts at most half Us length of the capsule they subtend — S . ravenii 64'. Bracts more than half the len e capsule they subten 65. Upper — erect 8 c. O. stricta subsp. argent inae 65’. Upper bracts — ud 66. Rosette leav cm wide; alins leaves 0.6- wide; bracts 0.7-1.2 cm wide; seeds 1.5-1.8 mm long Series I. RENNERIA 23a. O. stricta subsp. : stricta 66’. Rosette leaves 1.2— 1.2-2 cm wide; seeds 1.2-1.5 mm long . 28c. O. hus ap subs. brasiliensis Oenothera sect. Oenothera subsect. Munzia series Renneria (Fischer) Diet- rich, stat. nov. Based on Oenothera subgen. Renneria Fischer, Feddes Repert. Spec. Nov. Regni Veg. 64: 233. 1962 Onagra sensu Krause, Repert. Spec. Nov. Regni Veg. 1: 167. 1905. Oenothera $ Euoenothera sensu Munz & Johnston, I Gray Herb. 75: 17. 1925, pro parte. Oenothera subgen. Euoenothera sensu Munz, Physis 11: 280. ee 2 1933; Revista Univ. (Santiago ) 1937, pro parte; sensu Hagen, EE 1185 Publ. Sci. Ser. 16: 306. 1950, pro Oenothera subgen. Raimannia Munz, Physis 11: 279. 1933, pro parte; Amer. J. Bot. 22: 645. , pro parte 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 451 Erect annual or biennial, possibly sometimes perennial, herbs, forming a rosette or elongating soon after germination, sometimes flowering in the rosette, simple or with a + branched main stem and arcuate to oblique side branches arising from the rosette; plants less than 1 dm to 15 dm tall. Plants very densely to sparsely strigillose, often also appressed or erect long-villous, rarely glandular- pubescent. Rosette leaves linear to lanceolate, acute, narrowed gradually or + abruptly to the petiole, rarely sessile, 2.5-35 cm long, 0.3-6 cm wide; cauline leaves linear, elliptic or lanceolate, acute, sessile and acute to truncate at the base, or narrowly cuneate at the base and with a long or short petiole, 3-20 cm long, 0.3-3 cm wide; bracts linear, elliptic or lanceolate, acute, sessile and acute to subcordate at the base or attenuate at the base and with a long or short petiole, 3-12 cm long, 0.3-2.5 cm wide, longer than the capsules they subtend; leaves + regularly or irregularly serrate, plane or undulate at the mar- gins, sometimes (in O. nana) flecked with black. Inflorescence simple or branched, mostly crowded; flowers mostly obliquely erect, (1-)2-6 opening toward the apex each day. Floral tube 0.5-10 cm long, usually curved. Buds oblong to narrowly ovate in outline, 0.4-3.5 cm long, 3-8 mm thick, often with red stripes at the junction of the sepals with the floral tube, and sometimes also along the midveins. Sepals green, yellowish, or yellowish green, often flushed with red; apices of the sepals 14 mm long, erect or divergent. Petals very broadly obovate, rounded or retuse, 0.5-4 cm long, yellow or bright yellow, sometimes flushed with red along the veins and at the base, or completely red. Ovary 0.8-1 cm long. Capsule oblong or lanceolate to ovate in outline, 1-3 cm long, 3-10 mm thick, erect and + appressed to the stem, or (in O. nana) stand- ing out at right angles from the prostrate stems, evidently fused with the bracts: valves spreading apart in dehiscence or slightly incurved at the apex only. Seeds + obtusely angular, rounded and elliptic in outline, 0.8-1.7 mm long, 0.5-1 mm thick, light brown to dark brown or black, sometimes with darker flecks. Self- compatible; outcrossing or self-pollinating chromosomal homozygotes or self- pollinating complex heterozygotes. Gametic chromosome number, n = 7 (bi- valents, ring of 14, or intermediate configurations at meiotic metaphase I). Lectotype species: Oenothera versicolor Lehm: = O. campylocalyx Koch & Bouché. Distribution (Fig. 7): All species of this subsection are exclusively moun- tain plants and occur at elevations from 1,500 to 5,000 m, rarely lower. They oc- cur at these elevations in the Cordillera Occidental and Cordillera Oriental of Peru from the province of Lambeyeque southward a short distance over the Chilean border. In the Bolivian and Argentine Andes, the group ranges along the eastern flanks southward to Mendoza. Disjunct stations occur in the Sierra of Cordoba and in the Andes of Ecuador near Quito, the latter probably resulting from the naturalization of cultivated plants. The characteristic aspect of the plants of this section results from their thick stems, sometimes up to 2 cm in diameter, crowded inflorescences, and obliquely ascending flowers with somewhat curved floral tubes. Most diagnostic, owever, are the erect, urn-shaped or broadly cylindrical capsules (Fig. 9). The 452 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 — o G G SO (9 3-24. Leaves of South American taxa of Oenothera sect. Oenothera.—13. sette leaf of O. peruana (Peru, Arequipa, Santarius 2090 ).—14-16. Rosette leaf, cauline leaf, and bract of O. versicolor ( Bolivia, Cochabamba, Diers in 1959).—17-19. Rosette leaf, cau- line leaf, and bract of O. tafiensis subsp. tafiensis ( Argentina, Tucumán, Santarius 1733).— 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 453 three first species are annual, although potentially overwintering as a rosette and therefore biennial, whereas O. santarii and O. longituba are obligate biennials in nature, although flowering the first year in cultivation. These five species have somewhat angular seeds which are dark brown or black. Oenothera scabra forms only a few leaves near the base on germination, and then grows into a branching plant without forming a rosette. Its seeds are rounded in outline, smaller than those of the first-listed and presumably more generalized species of the series, light brown, and flecked. Oenothera tafiensis and O. pedunculifolia can be considered intermediate between O. scabra and the remaining species; thus O. tafiensis 77 155 tafiensis resembles O. peruana in habit, but has small flecked seeds. In O. tafiensis subsp. parviflora and O. pedunculifolia, both also with flecked seeds, the plants bolt from a rosette of limited size and resemble O. scabra in habit. 1. Oenothera peruana Dietrich, sp. nov.—Fics. 13, 102, 168. Herba annua vel biennis, erecta, rosulata, 5-10 dm alta, simplex vel caulis principalis ramosus et ramis oblique e rosula f ascendentibus. Herba tota dense strigulosa, circum inflores- centiam pilis 5 rahe praedita. Folia rosulae angustissime elliptica vel anguste oblance- olata, acuta, lamina in petiolum brevem gradatim decrescens, I —20 cm longa, 2—5 cm lata; folia caulina 8 vel cultrata ad anguste lanceolata, acuta, basi acuta, 6-20 cm longa, 1-3 cm lata, breviter [uq bractea l ia vel anguste “bivio acuta, basi truncata, ses- silia, 3-6 cm longa, 0.8-2 cm lata; folia omnia irregulariter + serrata. Eiflorescentia igs lex. ice floralis 1. 2-9 cm E. Gemmae ambito lanceolatae, 1.5-2.5 cm longae, 5-7 mm cras- , griseovirides vel flavovirides; apices sepalorum ca. 2 mm longi, erecti. Petala latissime un floride lutea, post anthesem decolorata, 1.5-2.5 cm longa. Stylus longus, stigmate sub anthesi supra antheras elevato. Gann 7-10 mm longum. Capsula 2-3 cm longa, qs mm crassa, maturitate brunnea. Semina obtuse angulata, 1.2-1.7 mm longa, 1-1.2 mm crassa, Numerus gameticus chromosomaticus, n — 7; planta chromosomatice admodem Eu c. Erect annual or biennial herb, forming a rosette, 5-10 dm tall. Main stem simple or somewhat branched, with side branches arising obliquely from the rosette. Entire plant densely strigillose, in the region of the inflorescence also with appressed long-villous pubescence. Rosette leaves very narrowly elliptic to narrowly oblanceolate, acute, gradually narrowed to the petiole, 15-20 cm long, 2-5 cm wide; cauline leaves linear or cultrate to narrowly lanceolate, acute, acute at the base, 6-20 cm long, 1-3 cm wide, with a short petiole; bracts linear to narrowly obovate, acute, truncate at the base, sessile, 3-6 cm long, 0.8-2 cm wide; all leaves irregularly and = prominently serrate, plane or undu- late at the margins. Inflorescence unbranched. Floral tube 1.2-2 cm long. Buds lanceolate in outline, 1.5-2.5 cm long, 5-7 mm thick, gray green or yellowish green; apices of sepals erect, ca. 2 mm long. Petals very broadly obovate, bright yellow, fading after anthesis, 1.5-2.5 cm long. Anthers 6-9 mm long. Filaments 8-11 mm long. Style long, the style held above the anthers at anthesis, 24.5 cm long. Stigma lobes 4-7 mm long. Ovary 7-10 mm long. Capsule 2-3 cm long, < 20-22. Rosette leaf, cauline leaf, and bract of O. santarii (Argentina, Mendoza, Santarius 1430 ).—23. Rosette leaf of O. tafiensis subsp. parviflora (Argentina, Tucumán, Göpel in 1961 ).—24. Rosette leaf of O. nana (Peru, Puno, Santarius 2045). 454 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 6-9 mm thick, dark brown to very dark brown when ripe. Seeds 1.2-1.7 mm long, 1-1.2 mm thick, obtusely angular, black. Self-compatible but outcrossing. Gametic chromosome number, n — 7 (7 bivalents*, ring of 8 and 3 bivalents **, ring of 12 and 1 bivalent ***, or ring of 14**** at meiotic metaphase I). Flow- ering time: December-April. Type: Grown from seeds and cultivated in the Botanical Garden of Düssel- dorf, Germany, 25 July 1972. Source: Peru, Dep. Arequipa, moist slope and edge of an irrigation ditch 19 km behind Arequipa at road to Chihuata/Puno, 2,700 m, 13 Apr. 1968, K. A. Santarius 2090 (MO-2155406, holotype; CTES, DUSS, M, isotypes). Distribution (Fig. 225): Very local in the Andes of south Peru and north Chile, 2,500-3,300 m elevation. Specimens examined a cultivated plants: Peru. AREQUIPA: Ca. 19 km 7 Arequipa at the road to Chihuata/Puno, 2,700 m, Santarius 2078. 2088 *. 5 * 2096***. 2098*, 2102“, 2103“, 2104 **, 27067, 2107*, ** 21 08** ** (DUSS, nios 1 CTES, 2075, 2088, 2090, 2096, 2098, 2106, 2108 also MO; 2090, 2098, 2106 also N ous specimens examin ie ERU. APURIMAC: Abancay, 3,100 m, Iltis & Ugent 695 (UC). AREQUIPA: Chihuata, 20 km E of Arequipa, Munz 15539 B POM). Pichu-Pichu, 3,300 m, Sandemann 3778 (K). Cai- lloma, 2 a m, Lopez 284 (RSA E. TARAPACA: Mamina, 8 9451 (CONC). In contrast to all of the following species, O. peruana has clearly angled seeds. On the basis of this characteristic, it seems to stand closest to the common origin of the South American species of sect. Oenothera with the North Ameri- can ones that have been called subgen. Euoenothera. It is possible that complex heterozygotes arise from time to time in the zone of overlap between O. peruana and O. versicolor, since some plants with a ring of 14 (Santarius 2108) are intermediate between these species. These plants are morphologically indistinguishable from O. peruana, and are not taken as representing an independent species. Plants with smaller rings of chromosomes are regarded as hybrids between chromosomally structurally differentiated indi- viduals of O. peruana. 2. Oenothera versicolor Lehm., Ind. Sem. Hort. Bot. Hamb. 7. Dec. 1855; Linnaea 28: 359. 1856.—Fics. 14-16, 103-104, 169. O. campylocalyx Koch & Bouché, Ind. Sem. Hort. Berol. App. 17. 1856. LecroryPE: Culti- Berlin Botanical Garden, 1832, ex herb. Kunth, source unknown ( B, destroyed; POM, fragments and 5 phs). O. coccinea Britton, Bull. Torrey Bot. Club 17: 213. 1890. rrEcrorvPr: Bolivia, Dep. Potosi, Ingenio del Oro, 3,280 m, Mar. 1886, H. H. Rusby 1815 (NY); Munz & Johnston, Contr. erb. 75: 22. 19 Onagra fusca Krause, Repert. Spec; Nov. Regni Veg. 1: 167. 1905. TYPE: Peru, Dep. An- cash, near Pampa Romas, between Samanco and Caraz, among herbs, especially grasses and shrubs, 3,200-3,500 m, A. Weberbauer 3211 (B, destroyed in World War II, POM, ae zraph Oe inis campylocalyx sensu Macbride, Field Mus. Nat. Hist., Bot. Ser., 13(4): 535. 1941, O. 557 sensu Macbride, Field Mus. Nat. Hist., Bot. Ser., 13 (4): 540. 1941, pro pa O. kopenhagensis Latzel, Biol. Zentralbl. 86. 409. 1967. Invalid name. = = Op Sy ii versi- color Lehm O. 5 sensu Munz, Opera Bot., Ser. B, 3: 39. 1974, pro parte. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 455 Erect annual or biennial herb, forming a rosette, 5-8 dm tall. Plants un- branched or with a scarcely branched main stem and with arcuate- ascending side branches from the rosette. Lower portion of plants sparsely strigillose, becoming densely so in the region of the inflorescence, where also with an admixture of appressed and erect long-villous pubescence. Rosette leaves nar- rowly lanceolate, acute, gradually narrowed to the petiole, 20-25 cm long, 1.5- 3.9 cm wide; cauline leaves very narrowly elliptic to narrowly lanceolate, acute, narrowly cuneate at the base, 8-12 cm long, 0.6-2 cm wide, sessile or with a short petiole; bracts linear to very narrowly elliptic or narrowly lanceolate, acute, acute to truncate at the base, 5-10 cm long, 0.5-2 cm wide; leaves plane or undulate at the margins, + regularly serrate with acute or rounded teeth, the margins reddish, especially in the bracts of the young buds. Inflores- cence mostly unbranched. Floral tube 1.2-2.5 cm long. Buds lanceolate in out- line, 1.5-2 cm long, 5-7 mm thick when mature, reddish at the junction of the sepals with the floral tube; apices of the sepals erect or divergent, 2-3 mm long. Petals very broadly obovate, rounded or retuse, 1.2-2 cm long, yellow, red near the base and along the veins or entirely red. Anthers 5-8 mm long. Filaments 6-10 mm long. Style mostly short, the anthers shedding pollen directly on the stigma at anthesis, but sometimes longer, the stigma elevated above the anthers, 2-3.5 cm long. Stigma lobes 2.5-4.5 mm long. Ovary 8-10 mm long. Capsule 1.5-3 cm long, 5-9 mm thick, glabrescent, dark brown to very dark brown when ripe. Seeds 1-1.3 mm long, 0.6-0.7 mm thick, + obscurely angled, dark brown to black. Self- oue self-pollinating or outcrossing. Gametic chromosome number, n = 7 (7 bivalents* at meiotic metaphase I). Flowering time: Ecua- dor and Peru, be lee Bolivia, October-May; Argentina, October-April. Type: not seen. The original description clearly indicates the species de- scribed here. Distribution (Fig. 226): Andes of Peru, Bolivia, and Argentina from the department of Ancash in Peru through Bolivia to the province of La Rioja in Argentina, 2,000-4,500 m elevation. The occurrence in Ecuador ( Pinchincha Prov.) might represent the establishment of adventive plants. Specimens examined from 5 plant Peru. Junin: Valley of Rio Mantaro, Pee: 3,650 m, Santarius 2163*, 2180, 2184 (DUSS, MO; 2163 also CTES, M). puso: Chimu, 8 km SE of Pans , Santarius 2062* (DUSS). BOLIVIA. COCHABAMBA: Tiy riuni near Cochabamba. Diers in April 1959* (CTES, DUSS, MO). rariya: Road from Tarija to Villazon, km 33, 3,100 m, Santarius 1946* (CTES, DUSS, M, MO). Additional specimens examined: NCHINCHA; Between Nono and Cotocollao, 3,100 m, Asplund 20276 (L NY, R, S, UPS). W of Eos 2,700 m, Harling et al. 10253 (RSA). PERU. ANCASH: Hua las, Laguna Paron, 3,000 m, Lopez 1878 (R . HUÁNUCO: Huacachi, near Muña, Macbride 3884 (F, G, GH, US). Pillao, 2,700 m, Wont in 1946 (F, MO, LIL, UC). uma: Rio Blanco, 3.900 m, Macbride > Featherstone 681 (F, GH). Rimac valley, 3,500 m, N.N. in 1954 (R SA). Chiela near Huarochiri, 4,000 m, Sanders 594 . Oroya near Line, Kalenborn 41 (MO), 241 (GH, NY, US). AYACUCHO: mpa de Chupas, N.N. in 1969 (RSA). APURIMAC: Apurimac River near Cañas, 3,700 m, Vargas 11044 (F, GH, K, UC). cuzco: Cuzco, Herrera in 1922 (SI); 3,350 m, Balfour 102 (K). Paucar- tambo, Hone cilia: 2,200 m, Vargas 9976 (MO). Ollantaytambo 3,000 m, Cook & Gilbert 628 (US). PUNO: Puno, 4,000 m, Soukup 109 (F). Carabaya, Antapampa, 4,180 m, Vargas — 456 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficures 25-34. Leaves of South S Esa taxa of Oenothera sect. Oenothera (contin- ued).—25. Rosette leaf of O. longituba (Argentina, Catamarca, Diers in 1959).—26. Bract O. tarijensis (Bolivia, Tarija Santarius 1924).—27. Bract of O. recurva (Bolivia, Tarija, Santarius 1941).—28. Bract of O. scabra (Bolivia, Cochabamba, Santarius 2003 ).—29-31. Rosette leaf, cauline leaf, and bract of O. longituba (Argentina, Jujuy, Fabris 5787 ).—32-34. Rosette leaf, cauline leaf, and bract of O. pedunculifolia (Argentina, Tucuman, Santarius 1745). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 457 7002 pro parte (LIL, POM). Azángaro, Tequena near Arapa, 3,900 m, Aguilar 227 (USM). Capachica Peninsula of Titicaca Lake, 3,700 m, Tutin 1009 (B OLIVIA. LA PAZ: La Paz, 3,800 m, Buchtien 532 (GH, NY, UC , US). Tacacoma, Cár- denas 5137 (US). Charasani near Muñecas, =. 700 m, Cardenas 3844 (POM). Quime, 2,400 m, Brooke 5396 (BM). Isla Titicaca, 3.840 n Buchtien 2921 (NY, US). Pongo, 3,650 m, Tate 190 (NY). Titicaca, Adolph & Bandelier 3 1905 (NY). Vicinity of Sorata near Lare- caja, 3,200 m, Mandon 628 (P pro parte, S), 627 (BM, G, K, P, W pro parte). Gran Poder near Sorata, 3, 050 m, Brooke 6644 (BM). cocHABAMBA: 30 mi ENE Cochabamba, 3,750 m, Brooke 5091 (BM). Choro above the 5 River, nd m, Brooke 6028 (BM, F, NY). ORURO: Caranga, D'Orbigny in 1929 (P). porosi: Ingen o de Oro o, 3,050 m, Rusby 1976 (NY). CHUQUISACA: Cordillera de E ae 3.100 m, Troll 670 (M). TARIJA: La 3 near Tarija, 3,300 m, Bolts 6112 (K, UC, US). ARGENTINA. JUJUY: Chorru Valley near Tilcara, 3,400 m, Balls 6027 E Mad Falda Grande, Cerro de G T ee 3,400 m, Cabrera & Hernandez 14021 (LP). a de Zenta, 4,000—4,500 m, Budín 178 (GH, POM). Tumbaya, Medinacely 9 (POM E ‘Cerro Morado near Tumbaya, 3,300 m, Fabris 6217 (BAA); Cabrera 18306 (LP). Volcan near Tumbaya, 2,900 m, Sleumer 3563 (LIL). Serrania de Calilegua, Tolditos, 2,600 m, Fabris et al. (LP) Serrania de Calilegua, Cerro Colorado, 2,700 m, Fabris et al. (LIL, LP). 5 Tascal near Tumbaya, 3,400 m, Cabrera 15105 (LP). Puente del Diablo near Tres Cruces, 4, ,000 m, Fabris & Marchionni 1746 (LP). Mina Aguilar near Tumahuaca, Fernández 2003 (LP). Mina Aguilar, 4,400 m, Sleumer 3383 (LIL). Mula Muerte, 2,700 m, Castillón 119, 130 (LIL). SALTA: El Alisal near Cafayate, 2,800 m, in 1914 (L. IL). Santa viaa , 2,385 m, Sleumer 3780 (LIL). Lizoite near Sta. Victoria, 3,340 m, Meyer & Bianchi in 1940 (LIL). TUCUMÁN: La Ciénaga, Lorentz d» Hieronymus 687, 689 (CORD. GOET); Lillo 3707 (LIL). Cerro 3,000 m, Burkart 5366 (SI). Estancia Sta. Boza and Pto. La Cu 0 m, Venturi 3165 LIL, US). Malamala, 2,400 m, Lillo 3443 (LIL). El Chorro near "ERIS = ,500-3,000 m, Schraiter 1121 (LIL). caramarca: Andalgala, Jorgensen 1054 pro parte (LIL). LA RIOJA: Sierra de Famatima, slope of La Mesada, 3,500-3,700 m, Kurtz 13946 Eu» Between La Mesada and El Paso, 3,500-4,200 m, Kurtz 14040 (CORD). I Lg m from cultivated plants: Ex herb. J. Gay, Jardin Vilmorin, Paris, 17 Aug. 1859, (K; as O. versicolor). Hort. Paris, Aug. 1865, "B. Verloti (P; as O. campylocarpa ). The partial or pronounced red coloration of the veins and base of the petals and the predominantly strigillose pubescence are characteristic of O. versicolor. As might be expected from the very extensive range, there is a considerable and significant amount of variation, especially in the width of the leaves and their toothing. Plants intermediate with O. lasiocarpa occur in the provinces of Tucu- man and Jujuy in Argentina. It is not known whether these are spontaneous hybrids or complex heterozygotes which characterize entire populations. 3. Oenothera lasiocarpa Griseb., Abh. Kónigl. Ges. Wiss. Gottingen 19: 143. 1874.—Fics. 105, 170. O. campylocalyx sensu Munz, Physis 11: 286. 1933, pro parte. Erect annual or biennial herb, forming a rosette, 1-3.5 dm tall. Plants un- branched or with a somewhat branched main stem and widely arcuate-ascend- ing side branches arising from the rosette. Entire plant densely strigillose, especially in the region of the inflorescence, and also with erect long-villous pubescence. Rosette leaves narrowly oblanceolate to very narrowly elliptic, acute, gradually narrowed to the petiole, 6-12 cm long, 0.8-1.5 cm wide; cauline leaves mostly absent, since the plants generally bloom just above the rosette, but when present very narrowly elliptic, 5-10 cm long, with a short petiole or sessile; bracts narrowly lanceolate to oblanceolate, 3-8 cm long, 0.3-1.2 cm ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 — @ @® © (8) © © Ficures 35-57. Leaves of South American taxa of Oenothera sect. Oenothera (contin- ued).—35-36. Cauline leaf and bract of O. mendocinensis (Argentina, Buenos Aires, Santarius Rosette leaf and cauline leaf of O. odorata (Argentina, Chubut, Santarius 953). —40. Cauli nd bract of O. odorata (Argentina, Río Negro, Santarius 800).—41—43. Rodi leaf, cauline leaf, and bract of O. ravenii P ravenii (Brazil, Rio Grande do Sul, 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 459 wide, acute, acute to truncate at the base; leaves plane or undulate at the margins, irregularly serrate with coarse to slightly produced teeth. Inflorescence unbranched. Floral tube 1.5-2(-2.5) cm long. Buds oblong to narrowly ovate in outline, 1—2 cm long, 6-8 mm thick, often flushed with red; apices of the sepals erect, ca. 2 mm long. Petals very broadly obovate, 1.5-2 cm long, rounded or retuse, red near the base and along the veins. Anthers 4-6 mm long. Fila- ments 6-8 mm long. Style short, the stigma surrounded by the anthers at anthe- sis, or long, the stigma held above the anthers at anthesis, 2-3 cm long. Stigma lobes 3-5 mm long. Ovary 8-10 mm long. Capsule 1-2(-2.5) cm long, 5-8 mm thick, dark brown to very dark brown when ripe. Seeds 1-1.2 mm long, 0.6-0.8 mm thick, = obscurely angled, dark brown. Self-compatible; self-pollinating or outcrossing. Gametic chromosome number, n = 7 (7 bivalents* at meiotic meta- phase I). Flowering time: November-April. Type: Argentina, Prov. Catamarca, alpine meadows, Vayas Altas, Sierra de Belén, 2,950-3,650 m, mid-January 1872, T. G. Lorentz 633 (GOET, holotype; CORD, isotype). Distribution (Fig. 225): Andes of Argentina, in the provinces of Salta, Jujuy, Tucumán, Catamarca, La Rioja, and San Juan; 2,000-4,000 m elevation. Specimens examined from cultivated plants: ARGENTINA. TUCUMÁN: Tafi del Valle, 3,900 m, Diers in 1959* (CTES, DUSS, M, MO). Additional specimens examined: ONTINA: JUJUY: Quebrada near Susques, Cabrera 8760 (LP). Sierra de Zenta, 4,500 m, Budin 7476 (LIL). Yavi Chico, 3,350 m, Werner 286 (LP). Maimara, 2,230 m, Budin 11790 (GH, LIL, POM, SI). Volcán, Loma del Tambo, 2,500-3,000 m, Schreiter in ; L). SALTA: Vicinity of Nevado del Castillo, Lorentz & ieronymus 47 (CORD, ET). TucuMAN: Tafi, Cerro Muñoz, 3,000 m, Lillo 7428, 4267 (LIL); Castillon 177 (LIL). Rio Blanco, 2,600 m, Lillo 8868 (LIL). Lara near Tafi, 3,200 m, Rodriguez 574 (GH). Quebrada del Barón, Fabris 1371 (LP). Quebrada de la Hoyada, 1,500 m, Schreiter 1875 7 P). Peñas Azules, Parodi 10971 (POM); Burkart 5300 (SI); Olea 8759 (LIL). La Quenoa, 2,800 m, Schreiter 6995 (LIL). La Ciénaga, 2,800 m, Sleumer 203 (LIL). Pojonal, 2,600 m, Meyer 3659 (LIL). Casa de Piedra, Lillo 179 (LIL). Vicinity of Muñecas, Castillon in 1905 (LIL). Chicligasta, Estancia Las Pavas, 3,000 m, Venturi 4597 (BAB, LIL, SI). Tran- cas, Chorro, 3,400 m, Venturi 8512 (US); Schreiter 4790 (LIL). CATAMARCA: Andalgala, valley of Río Bolsón, 4,000 m, Rohmeder in 1943 (LIL). Campo Grande, Schickendantz 67 (CORD, GOET, SI); Schickendantz 310 (CORD, GOET). Sierra de Ambato, between El deo and Cerro Manchado, 2,900 m, Hunziker 19196, 20998 (CORD); Hunziker & Di Fulvio 19759, 19721 (CORD). Belén, slopes N of Portezuelo, 3,200-3,300 m, Sleumer d» Vervoorst 2606 (LIL, US). Tinogasta, La Tranca, Castellanos 564 (POM). La Puntilla near Tinogasta, 2,700 m, Hunziker 4356 (BAB). LA RIOJA: Sierra Famatima, La Vega de la Mesada, 3,650 m, Kurtz 13928 (CORD, MO). Ciénaga de La Caldera, 3,600-3,650 m, Kurtz 13935 (CORD). El Volcán, Kurtz 14658 (CORD). Mina San Juan, 3,050-3,200 m, Kurtz 13597, 13685 (CORD); Real Viejo, Kurtz 14701 (CORD). Alto Blanco, Castellanos 28-282 (POM). La ~ — Hackbart in 1966 ).—44—-46. Rosette leaf, cauline leaf, and bract of O. indecora subsp. bonari- ensis (Argentina, Corrientes, Quarín & Schinini 1297).—47-49, Rosette leaf, cauline leaf, and bract of O. ravenii subsp. chilensis (Chile, Cautin, Stubbe in 1960).—50-52. Rosette leaf, cauline leaf, and bract of O. longiflora subsp. grandiflora (Argentina, Corrientes, Krapovickas & Cristóbal 11293 ).—53-54. Cauline leaf and bract of O. catharinensis ( Brazil, Santa Cata- rina, Conrad & Dietrich 9).—55-57. Rosette leaf, cauline leaf, and bract of O. longiflora subsp. longiflora ( Uruguay, Colonia, Santarius 73). 460 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 h ) @ © © Ficures 58-82. Leaves of South American taxa of Oenothera sect. Oenothera (continued). —58-59. Cauline leaf and bract of O. affinis gentina, San Luis, Conrad & Dietrich 139). —60-61. Cauline leaf and bract of O. mollissima (Uruguay, Maldonado, Santarius 161 62. Bract of O. mollissima (Uruguay, Maldonado, Santarius 162).—63-64, Cauline leaf d > " = 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 461 Mejicana, 3,900 m, Parodi 7896 (GH), Quebrada de Potrerillos, 3,800 m, Krapovickas & Hunziker 5411, 5419 (BAB). Cueva de Perez, Hieronymus & Niederlein 390 (CORD). Mina Oro, Hieronymus & Niederlein 414 (CORD). Lamadrid, Realitos, 3,800 m, Rohmeder in 1941 (LIL). Real de los Neveros, 3,500 m (LIL-80298). Viejo, Hossens 1422 (CORD). san JUAN: Cordillera " Salado, 3,800 m, Spegazzini in 1957 (BAB). Cerro Tronador, Spegazzini 212 pro parte (BAB). Oenothera lasiocarpa is very closely related to O. versicolor, and the two are the only species of the series with pronounced red coloration of the petals. It can be separated from that species by its lower and more compact habit, erect long-villous pubescence, and dark green leaves; in O. versicolor the leaves are more grayish green. Small plants of O. lasiocarpa can also be confused with O. nana, but the latter does not develop a central shoot at all. Very small-leaved plants occur in O. lasiocarpa (e.g., Budín 1170), and some plants of O. nana growing in the area of O. lasiocarpa have broad leaves (Lillo 5516, Rodríguez 1292, de la Sota 2711) which, despite their occurrence as rosettes, resemble O. lasiocarpa closely. The first group of plants may be understood as occasional homozygous derivatives of O. nana. Similarly, the broad-leaved plants of O. nana may provide an indication that O. lasiocarpa, which is in all probability derived from O. versicolor in Argentina, may be involved in the origin of one of the chromosomal complexes of O. nana, which is always a complex hetero- zygote. 4. Oenothera santarii Dietrich, sp. nov.—Fics. 5, 20-22, 106, 171, 195. O. odorata Jacq, var. brachycarpa Haumann, Anales Soc. Ci. Argent. 86: 292. 1919. LECTO- TYI entina, Prov. Mendoza, Cordilleras Altas de Mendoza, Puente del Inca to Punta de | c ca. 2,500 m, Jan. 1908, L. Haumann (BR). erba annua vel biennis, rosulata, 7-12 dm alta, simplex vel caulis principalis vix ramosa, ; rescen petiolum duas decrescens, 15-30 cm ene 25-4 cm m r folia caulina angustissime ellip- tica, acuta, basi acuta, 10-15 cm longa, 1-2.5 cm lata, brevipetiolata vel sessilia; bractea 5 elliptica vel lanceolata, basi attenuata vel truncata, plerumque o lique ascenden- tia, sessilia, 5-8 cm longa, 1-1.5 cm lata, apicibus supremarum saepe in cochleam tortis; folia valde marginibus undulatis. Inflorescentia plerumque simplex. Tubus floralis 2-2.5 cm lon- us. Gemmae ambito lanceolatae, 2-2.5 cm longae, 5-8 mm crassae; apices sepalorum ca. 2 ë , 8-10 - mm A saan Capsula 23 cm longa, 6-9 mm crassa, maturitate fusca. Semina + obtuse angulata, 1.2-1.4 mm longa, 0.7-1 mm lata, fusca. Numerus gameticus chromosomaticus, n = 7; planta chromosomatice homozygotica < bract of O. rivadaviae (Argentina, Chubut, Santarius 913).—65-66. Cauline leaf and bract of O. picensis subsp. cordobensis (Argentina, Córdoba, Gópel in 1961 ).—67-69. Rosette leaf, cauline leaf, and bract of O. stricta subsp. sie (Argentina, Rio Negro, Santarius 798 ).— 70-72 sette leaf, cauline leaf, and bract of O. stricta subsp un te (Arg a, Buenos Aires, Santarius 346).—73-75. Rosette cauline leaf, ct of O. bahia-blancae (Ar- entina, Buenos Aires, Santarius 455).—76-78. Rosette leaf, cauline leaf, and bract of O parodiana subsp. parodiana (Argentina, Cór , Conrad e Dietrich 98).—79-81. Rosette leaf, cauline leaf, and bract of O. parodiana sub Jap. parodiana (Argentina, Córdoba, Conrad 2 Dietrich 162). —82. Rosette leaf of O. parodiana subsp. parodiana (Argentina, Córdoba, Conrad & Dietrich 122). 462 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Erect annual or biennial herbs, forming a rosette, 7-12 dm tall. Plants unbranched or with a scarcely branched main stem and widely arcuate-ascend- ing side branches arising from the rosette. Plant sparsely strigillose, somewhat more densely so in the inflorescence, also with sparse, erect, villous pubescence. Rosette leaves narrowly oblanceolate, acute, gradually narrowed to the petiole, 15-30 cm long, 2.5-4 cm wide; cauline leaves very narrowly elliptic, acute, acute at the base, 10-15 cm long, 1-2.5 cm wide, with a short petiole or sessile; bracts very narrowly elliptic to lanceolate, acute, acute to truncate at the base, usually obliquely ascending, sessile, 5-8 cm long, 1-1.5 cm wide, the tips of the uppermost ones often slightly spirally twisted; all leaves plane to strongly undulate at the margins and irregularly toothed. Inflorescence usually unbranched. Floral tube 2-2.5 cm long. Buds lanceolate in outline, 2-2.5 cm long, 5-8 mm thick; apices of the sepals erect, ca. 2 mm long. Petals very broadly obovate, yellow, 2-3 cm long. Anthers 9-11 mm long. Filaments 15-20 mm long. Style long, the stigma elevated above the anthers at anthesis, or short, the anthers shedding pollen directly on the stigma at anthesis, 3-5.5 cm long. Stigma lobes 3-5 mm long. Ovary 8-10 mm long. Capsule 2-3 cm long, 6-9 mm thick, brown when ripe. Seeds 1.2-1.4 mm long, 0.7-1 mm thick, more or less obtusely angular, brown. Self-compatible. Cametic chromosome num- ber, n — 7 (7 bivalents* or small rings at meiotic metaphase I). Flowering time: November-March. Type: Grown from seeds and cultivated at the Botanical Garden of Düssel- dorf, Germany, 15 Aug. 1972. Source: Argentina, Prov. Mendoza, Cordillera de Los Andes, on stony places at Ruta 7, ca. 150 m E of Arroyo Santa Maria, ca. 6 km E of Puente del Inca, 2,750 m, 25 Feb. 1968, K. A. Santarius 1430 (MO- 2155403, holotype; CTES, DUSS, M, isotypes). Distribution (Figs. 225, 241): Andes of Argentina west of Mendoza, and one locality in the province of San Juan, as well as in the Sierra de Córdoba and the Sierra de Comechingones (provinces of Córdoba and San Luis), 1,800-3,000 — m elevation. Specimens examined from cultivated plants: ARGENTINA: MENDOZA: Cordillera de los Andes, Puente del Inca, 2,700-2,750 m, San- tarius 1405* 1417 *, 1430*, 1436*, 1443 * 1444*, 1455 (DUSS; 1430, 1435, 1436, 1455 also M; 1430 1444 also MO). Punta de Vacas, 2.450 a cd 1492, 1500* (DUSS; 1492 also MO). aaa 2,450 m, sr 1535 (DUSS, MO). san Luis: Sierra de Come- chingones, El Rincón, 1,500 m, Conrad & Dietrich T (ring of 4 and 5 bivalents, ring of 6 and 4 5 8). ddit S = iv] 3 A N [Ke] — ws — — O > — E Z — > > = te The species, which has been known for a long time in the literature and in the herbarium as O. nana, O. punae, or O. kuntziana, is made up of two differ- ent elements. To the first belong plants with two chromosomal complexes from series Renneria, and this group includes the type of O. nana. To the second group belong plants in which are combined one genome from series Renneria and one from series Allochroa. These plants belong by definition to subsect. Clelandia. For them, the name O. punae is to be used. It is clear that the complex heterozygous O. nana, with its very condensed habit, has been derived from plants of normal stature in the process of adapta- tion to the extreme conditions at the high elevations where it occurs. I have already referred to O. lasiocarpa as a transitional species, and there are plants in O. nana which have some of the characteristics of that species. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 489 On account of the extremely condensed habit of O. nana, a complex mode of adaptation that alters the expression of many characteristics, it is difficult to analyze the origin of this species. The analysis of hybrids with species of nor- mal stature has revealed, however, the probable influence of O. peruana, O. versicolor, O. lasiocarpa, and O. scabra each with varying expression of their features in O. nana. The influence of O. versicolor and O. lasiocarpa appears to be stronger in the southern part of the range of O. nana, and that of O. peruana and O. scabra seems stronger in the north. The individuals in a population are rarely uniform, but vary in the same way as has been discussed for O. sandiana. The situation is made still more complex by the fact that O. nana occurs together with O. punae at many localities, and their chromosomal complexes are essentially interchangeable with one another; see also the remarks on p. 611 in this connection. The nanism of O. nana, its most prominent characteristic, can be manifested in various ways. Both complexes can have the dominant traits for normal size expressed, or one complex can be dominant and the other intermediate or recessive. Series II. ALLOCHROA Oenothera sect. Oenothera subsect. Munzia series Allochroa (Fischer & Meyer) Dietrich, comb. nov. Based on Oenothera sect. Allochroa Fischer & Meyer, Ind. Sem. Hort. Petrop. 2: 44. 1836 Onagra sensu Moench, Meth. Pl. 1: 675. 1794, pro parte; Suppl. Meth. Pl. 2: 287. 1802, O parte. Oenothera sect. Onagra Séringe ex DC., Prodr. 4: 46, 1828, pro parte. Oenothera sensu Spach, Nouv. Ann. Mus. Hist. Nat. 341. 1835, pro parte. Oenothera sensu Raimann, in Engler & Prantl, Nat. Pflanzenfam. III, 7: 214. ra pro parte. idi is sensu Sprague & Riley, irs Misc. Infor. 1921: 200. 1921, pro part Oenothera $ Raimannia sensu Munz & Johnston, conte. Gray Herb. 75: 16. 1985, pro parte. ipt iud subgen. Raimannia Munz, Physis 11: i . 1933, pro parte; Amer. J. B 45. 35, pro parte; Revista Univ. (Santiago) 2 22: 261. 1937, pro parte; nun: Bot. Mus. Hist. Nat. Montevideo 1(10): 26. 1943. Oenothera subgen. Raimannia sect. Raimannia Munz, North Amer. Fl., ser. 2, 5: 105. 1965, O parte. ot. 22: Erect annual or biennial herbs, rarely prostrate, forming a rosette or the stem elongating soon after the development of a few basal leaves, unbranched or with a branched main stem and ascending branches from the rosette which either arch outward in ascending or rise sharply and abruptly; plants 0.5-15 dm tall, rarely even taller. Stems more slender than in series Renneria, 2 to at most 10 mm thick. Plants (1) exclusively strigillose; (2) densely to sparsely strigil- lose, densely to sparsely long- and short-villous, the hairs mostly appressed, and densely to sparsely glandular-pubescent; (3) very densely to sparsely long- and short-villous and densely to sparsely glandular-pubescent; or (4) densely short- villous and densely glandular-pubescent. Rosette leaves linear to oblong, very narrowly oblanceolate to oblanceolate or narrowly elliptic, long- or short-acute, sessile and narrowly cuneate to truncate at the base or gradually narrowed to the petiole, 8-25 cm long, 0.4-3.5 cm wide; cauline leaves linear to oblong, very 490 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 narrowly elliptic to elliptic or narrowly lanceolate to narrowly ovate, acute, sessile, narrowly cuneate to truncate at the base, 2-20 cm long, 0.2-3 cm wide; bracts linear to broadly oblong, narrowly elliptic to elliptic or narrowly lanceo- late to ovate, acute to obtuse, rarely almost rounded, sessile, acute to sub- cordate at the base, 2-10 cm long, 0.2-2.5 cm wide; leaves mostly irregularly and distantly serrate with dull teeth, occasionally very coarsely serrate or doubly serrate (O. coquimbensis), plane or evidently to very slightly undulate along the edges. Inflorescence unbranched or branched, lax; flowers erect in anthesis, one opening each day. Floral tube 0.5-13 cm long. Buds oblong, elliptic to broadly elliptic or narrowly lanceolate to narrowly ovate in outline, 0.3-3.5 cm long, 2-11 mm thick, often with red stripes at the junction with the floral tube. Sepals green or yellowish green, often flushed with red, sometimes densely to sparsely flecked with dark red; apices of the sepals 0.5-4 mm long, erect, spreading, or hornlike. Petals obovate to very broadly obovate, rarely elliptic to broadly elliptic, rounded or retuse, 0.3-5 cm long, yellow to bright yellow, sometimes with a red basal spot. Ovary 1-2.5 cm long. Capsule linear to narrowly oblong in outline, terete, rarely slightly enlarged in the upper third or tapering at both ends and appearing short-petiolate, projecting obliquely from the stem, straight or slightly curved, (1.5-)2-6 cm long, 2-5 mm thick, not fused with the bract; valves curving outward or inward after the capsule dehisces, occasionally spreading. Seeds 1-2 mm long, 0.4-1.1 mm thick, nar- rowly elliptic to rotund in outline, light or dark brown to almost black. Self- compatible; chromosomal homozygotes or self-pollinating complex heterozy- gotes, rarely outcrossing. Gametic chromosome number, n — 7 (7 bivalents, ring of 14 or intermediate configurations at meiotic metaphase I). e species: Oenothera mollissima L. Distribution (Fig. 7): These are predominantly plants of relatively low eleva- tions from sea level upward; ascending to 3,200 m elevation in the Andes (O. affinis, O. arequipensis, O. featherstonei, O. nocturna, O. odorata, O. verrucosa). In Brazil, the plants occur in Guanabara, southern Minas Gerais, and in Sao Paulo to Rio Grande do Sul. They occur throughout Uruguay and in all prov- inces of Argentina as far as Rio Gallegos in Patagonia, extending northward to Tarija in Bolivia. West of the Andes, they range in Chile from the provinces of Atacama to Magellanes, and in the coastal deserts and semideserts from the department of La Libertad in Peru to Valparaiso in northern Chile. In Peru they ascend into the mountains along river valleys to 3,200 m elevation. If the isolated stations that occur up to 3,200 m elevation in the Andes are disregarded, most species of this series are inhabitants of the broad plains of Argentina and the coastal regions on both the Atlantic and Pacific shores of southern South America, and the lowermost slopes of the mountains. All species have more slender stems and are more graceful than the sturdy, thick-stemmed species of series Renneria. Important differences from that series are the laxer inflorescences, erect flowers, and obliquely divergent capsules. Most of the species of series Allochroa form a rosette, like the majority of those of series Renneria. Among the chromosomally homozygous species, only DIETRICH—SOUTH AMERICAN OENOTHERA O. indecora Aires peni 978). —1. 3. O. affinis ( Argentina, O. mollissima ( Uruguay, Montevideo, lodos 42). —135. ). rivadaviae Biwentino. Chubut, Santarius 924). 492 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 O. affinis, O. catharinensis, O. coquimbensis, and O. verrucosa form no rosette; whereas among the complex heterozygotes, no rosette is formed by those which incorporate a genome from O. affinis: namely, O. mollissima, O. montevidensis, and O. picensis. 14. Oenothera mendocinensis Gillies ex Hooker & Arnott, Bot. Misc. 3: 310. 1833.—Fics. 35-36, 121, 174, 210. O. odorata sensu Hicken, Physis 2: 110. 1916. Raimannia mendocinensis (Gillies ex Hooker & Arnott) Sprague & Riley, Bull. Misc. Infor. 1921: 201. 1921. Oenothera argentinae H. Lév. & Thell. var. camptotricha Kloos & Thell., Ned. Kruidk. Arch. 1921: 100. 1921. LEcrorvpE: Netherlands, Rotterdam, Maashaven meal factory, 10 Sep. 1920, A. W. Kloos (BAS). O. indecora sensu Munz, Physis 11: 281. 1933, pro parte; Amer. J. Bot. 22: 658. 1935, pro parte Erect annual herb, forming a rosette, unbranched or with a branched main stem and widely arching or obliquely ascending side branches arising from the rosette, 3-6 dm tall. Plants either exclusively and densely strigillose or densely to sparely strigillose and densely to sparely villous, but often strigillose only near the base. Rosette leaves linear, acute, narrowed to the petiole, 7-14 cm long, 3-5 mm wide; cauline leaves linear, acute, 3-6 cm long, 2-4 mm wide; bracts linear, acute, truncate at the base, sessile, 3-5 cm long, 1-3 mm wide, mostly somewhat longer than the capsules or + the same length; leaves plane or slightly undulate at the margins, irregularly serrate. Inflorescence branched. Floral tube 0.5-1.5 cm long. Buds oblong to elliptic in outline, gray green, often flushed with red, 4-9 mm long, 2-4 mm thick; apices of the sepals erect, 0.5-1 mm long. Petals very broadly obovate, 0.5-1 cm long. Anthers 2.5-5 mm long. Filaments 3-5 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 8-21 mm long. Stigma lobes 2-3.5 mm long. Ovary 1.2-1.8 cm long. Capsule (2-)3-6 cm long, 2-3 mm thick. Seeds elliptic in outline, light brown, (1-)1.2-1.8 mm long, 0.5-0.7 mm thick. Self-pollinating. Gametic chromosome number, n — 7 (7 bivalents* at meiotic metaphase I). Flowering time: October-March. Lectotype: Argentina, Prov. Mendoza, between Las Chacayes and Meloco- tón, foot of the Andes of Mendoza, (1824?), Gillies (K, POM photograph; a photograph of isolectotype at E, from Herb. GL). Two sheets of the type col- lection are in K, both mixed with O. indecora (one collected by Charles Darwin at Bahía Blanca in 1832). Because O. indecora does not occur in the province of Mendoza and because it cannot be determined with certainty which plants the labels correspond to, only one branch has been selected as lectotype. For this branch, the correspondence of the plant with the label is clear. Distribution (Fig. 226): Occurs only at low elevations, ascending to 1,500 m in the Andean foothills of Mendoza. The range includes the following prov- inces of Argentina: Mendoza, San Luis, Córdoba, Buenos Aires, La Pampa, Río Negro, Chubut, and Santa Cruz. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 493 Specimens examined from cultivated plants: ARGENTINA. BUENOS AIRES: Dunes ca. 2 km SE of Argerich, 37 km W of Bahia Blanca, Santarius 402*, 411*, 412, 415*, 417, 420*, 421*, 423, 425, 431*, 433, 435*, 438, 442*, 44 , 448 (DUSS; 402, 420. 421 alto CTES; 402, 415, 420, 421, 442 alse. M; 402, 420, 491, 425 also MO). Dunes along road from Villa del Mar to Ruta 229, ca. 5 km NNW of Punta ‘Alta, d 518*, 5921, 529, 533*, 536 are ses also M; 521, 533 also MO). CHUBUT: Dunes E of Puerto Madryn, Santarius 1354*, 1358, 1361, 1367, 1371, 1374*, 1378, 1382, 8 (DUSS; 1354, 1374 also CTES; 1 5 "ud 1374 alto M; 1354, 1371, 1374 also MO). 1 . examined: ARGENTINA. ENOS AIRES: Daireaux, Parodi 13163 (BAA). Monte Veloz, Parodi 12238 (BAA). "General "Villegas dunes near Bunge, Cabrera 5703 (LP). Between Piedritas and Canada Seca near General Villegas, Hunziker 12826 (CORD, MO). Salliquelo, Est. : Gorros” near Pellegrini, Cabrera 8012, (F, GH). Dunes near Pellegrini, Cabrera 6955 (LP). Pumta Alta, O'Donell 1513 (LIL). Colonel Dorrego, Monte Hermoso, Erettowi 2607 (MO). From Urdampilleta to mn on a 205 near Bolívar, Vervoorst 5437 (BAB). 25 km SE Carmen de Patagones, Fabr ofer 5001 (LP). Río Negro near Carmen de Patagones, Hunziker 414 (CORD); Mey er 6991 (LIL). Between Bahia San Blas and San Blas, Ame- ghino in 1903 (POM). Villarino near Bahía Blanca, Correa 2387 (BAB); Boelcke 11788 BAA, MO, SI). Ruta 22, Puerto 11552 (MVFA). Argerich near Villarino, Parodi 13799 (BAA, MO). Agustina near Junín, Cabrera 6560 (F, SP, NY). cónpoBA: Near Pacheco de Melo, road to Laboulaye, Hunziker 12768 ( CORD, MO), 12779 (CORD). Between Labou- lave and Salguero, Hunziker 12810 (CORD, MO, RSA). Sierra Chica, La Redución, Burkart 17325 (SI). Laguna Brava, 400 m, King 319 (BM). Quinta Soriano N of Bajo Grande near Córdoba, Kurtz 16128 (CORD, MO). Between Est. S. Miguel and Rufino near Santa Fé, Spegazzini 6915 pro parte (BAB). Between Rufino and La Cesira, kee 12853 (RSA). Sierra Ochoa, Stuckert 13562 (G). Dep. Unión, La Carlota, Hunziker 11206 (CORD, MO). Est. La Mascota near Ballesteros, Hunziker 12751 (CORD, RSA). SAN Luis: Laguna near Sayape, Castellanos in 1949 (LIL). Pedernera on Ruta 148, 10 km S Villa Mercedes, 510 m, Anderson 1338 (LP). Villa Mercedes, Corradi 4832 (SI). Between Villa Mercedes and Juan Jorba on ie 8, si o 13170 (CORD, cun Near Esquina on Ruta 7, between San Luis and E. Lo Tung nziker 13123 (CORD). Dep. General Pederrera, on Ruta 148 between Laraisse a El Durazno, 500 m, Hunziker 15977 (CORD). Nueva Escocia, Burkart 10840, 10808 (LIL, SI). LA PAMPA: Guatr aché, 98 8 94 (BAB). General Pico, Burkart 9919 (LIL, SI). General he Orbea in 1953 (SI, US); Burkart 19205 (SI). Between General Acha and Santa Ros Troncoso in 1959 (SI). General Lagos (SI-4856). Laguna La Asturi- ana, Bacigalupo in 1959 (P, SI). La Pampa, Monticelli 39 (SI). MENDOoZA: Las Heras, La Crucesita, Ruiz Leal 5378 (Leal). Mina Atalu, Ruiz Leal 3328 (Leal). Dep. Tunuyán, La Piedra Rajada, Ruiz Leal 1707 (Leal). Tupungato, road to Est. Silva, 1,500 m, Cáceres 6 (LIL, NY). 10 km W Campo de Los Andes, 1,500 m, Araque & Barkley 20Mz179 (LIL). Tupungato, Ruiz TU 2784a (LIL). Dep. San Carlos, Est. Viluco, Torres 36 (SI). Mendoza, Jörgensen 131 (BAB, C). Rio necro: S. Antonio, Guerin in 1910 (LIL). General — 250-360 m, Ficka 86 (BM, F, GH, K, MO, NY, SI, US). Dep. General Roca, banks of R Negro near J. J. Gomez, Krapovickas et al. (CTES). Río Negro, Berg 98 (CORD); N. N. 88 in 1874 (LE). SANTA CRUZ: 8 ory of 9 Cruz, Ameghino 32 (BA). Specimens from outside of South Americ NETHERLANDS. aaa 1902, Jansen p Wachter 13286 (L); 1931, Kern & Reichgelt 12181 (L). GERMANY. Emmerich on Rhine River, 1931, Kern & Reichgelt 5064 (L). Its small flowers, very small leaves, mostly long and narrow fruits, and strig- illose pubescence set off O. mendocinensis as an isolated species. On the basis of the pubescence, it might be regarded as the least specialized species of the series, since it is the only chromosomally homozygous member of series Allo- chroa which has retained this characteristic of the members of series Renneria. On account of its slender habit and small flowers, O. mendocinensis has been included incorrectly in the synonymy of O. indecora, on the assumption that a marked similarity between these species indicates a close relationship. The hybrid between Santarius 405 (O. mendocinensis) and O. argentinae "Erlangen" 494 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 (— O. indecora subsp. bonariensis) formed 7 bivalents at meiotic metaphase I, from which it cannot be inferred, however, that O. mendocinensis and O. in- decora are closely related, as discussed in the introduction. Oenothera mendocinensis is closely related to O. odorata, judged from their close similarity in habit, leaf form, and characteristics of the capsule and sceds. They have probably been derived from a common ancestor (Fig. 8), a sugges- tion which is consistent with their ranges. It appears likely that the more generalized ancestral forms in series Alloch- roa—the closest living equivalents of which may be O. mendocinensis and O. odorata—were probably derived from generalized members of series Renneria in a center which on the basis of present-day distributions would appear to have been in and about the province of Mendoza. They then migrated to the south and southeast. The more advanced species (O. indecora, O. ravenii, O. longiflora, O. affinis), on the other hand, may have originated farther north, in the region of the provinces of Catamarca and Tucumán, and then spread pre- dominantly toward the east and northeast and the region of the Chaco (Fig. 8). 15. Oenothera odorata Jacq., Icon. Pl. Rar. 3: tab. 456. 1795; Suppl. Coll. Bot. 5: 107. 1796.—Fics. 37-40, 122-123, 175, 211. Onagra undulata Moench, E Meth. Pl.: 287. 1802. type: The herbarium of Moench apparently no longer s. ido ire pris W. T. yee on, Hortus Kew. 2: 342. 1811. TYPE: Seeds from Port Desire, na, 1790, cultivated at Kew (not seen). O. oe var. virescens Séringe in DC., Prodr. 3: 48. 1828. Lecrorype: Hort. parisiensis, 6, M. Brun in 1824 (G-DC). Grown from seed from the same source as the type of ecies. odorata var. glaucescens Séringe in DC., Prodr. 3: 48. 1828. LECTOTYPE: 28 August, h.h. odoratissima Tausch, Flora 22: 557. 1839. LECTOTYPE: Cultivated in E ie garden, b. V Kosteletzky (PRC, POM photograph); Munz, Amer. J. Bot. 22: 661. 1935. E sensu Hooker & Arnott, Bot. Beech. Voy. 23. 1841. ibari Philippi, Anales Univ. Chile 84: 633. 1893. LECTOTYPE: Argentina, Prov. San Cruz, Lago Argentino ( Lago Santa Cruz), 30 Jan. 1879, E. Ibar (SGO, GH photograph, odorata f. glabrescens, media and undulata Spegazzini, Revista Fac. Agron. Univ. Nac Plata: 520. 1898. types: not locate . mollissima sensu Macloskie, Rep. Princeton Univ. Exped. Patagonia 8(5, 3): 613. 1905. Onothera polymorpha H. Lév. race odorata (Jacq.) H. Lév., Monogr. Onoth. 363. 1909; Bull. Acad. Int. Géogr. Bot. 19: 1909. O. 5 race odorata var. unata ( W. T. Aiton) H. Lév., Monogr. Onoth. 363. 1909; ull. Acad. Int. Géogr. Bot. 3. 1909. O; Pe a a race propinqua (Spach) H. Lév. var. ibari (Philippi) H. Lév., Monogr. Onoth. 365. Acad. Int. Géogr. Bot. 19: d mollissima subsp. odorata (]Jacq.) Thell. Mitt. Bot. Mus. Univ. Zürich 58: 390. S 99 ° ° ° ~ Raimannia odorata (Jacq.) Sprague & Riley, Bull. Misc. Infor. 1921: 201. 1921. Oenothera stricta sensu Munz, Physis 11: 285. 1933, pro parte; Amer. J. Bot. 22: 661. 1935, 0 EAR Erect annual herb, forming a rosette or the stem elongating after the forma- tion of a few basal leaves, unbranched or with a branched main stem and pros- trate or widely arcuate-spreading to obliquely ascending side branches arising from the rosette, 2.5-8 dm tall. Plants densely or sparsely strigillose and densely DIETRICH—SOUTH AMERICAN OENOTHERA IGUREs 136-139. Taxa of Oenothera sect. Oenothera subsect. Munzia.—136. O. stricta subsp. stricta (Chile, Cautin, Stubbe in 1960).—137. O. stricta subsp. altissima (Argentina, Rio Negro, Santarius 798 ).—138. O. stricta subsp. argentinae (Argentina, Buenos Aires, San- tarius 346).—139. O. bahia-blancae (Argent. na, Buenos Aires, Santarius 457) 496 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 or sparsely erect-villous or very densely to sparsely villous with erect long and short hairs, rarely sparsely glandular-pubescent. Rosette leaves linear to nar- rowly oblanceolate, acute, gradually narrowed to the petiole, 15-20 cm long, 0.5- 1.5 cm wide; cauline leaves linear to very narrowly elliptic or narrowly lanceolate, acute, narrowly cuneate to acute at the base, short-petiolate or sessile, 5-18 cm long, 0.5-1.5 cm wide; bracts narrowly lanceolate to narrowly ovate, acute, truncate to subcordate at the base, 3-7 cm long, (0.3-)0.5-1.5 cm wide, shorter than, equal to, or longer than the capsule; leaves plane or the margins markedly to slightly undulate, irregularly serrulate. Inflorescence unbranched. Floral tube (1.5-)2-3 cm long. Buds lanceolate to narrowly ovate in outline, green or yellowish green, often flushed with red, 2-3 cm long, 0.5-1 cm thick; apices of the sepals erect, sharply divergent, or hornlike, 2-3 mm long. Petals very broadly obovate, retuse, 24.5 cm long. Anthers 9-14 mm long. Filaments 16- 24 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, or long, the stigma held above the anthers at anthesis, 3.5-6.5 cm long. Stigma lobes 4-7 mm long. Ovary (1-)1.3-1.7 cm long. Capsule 3-5 cm long, mm thick. Seeds elliptic in outline, light brown, 1.5-2 mm long, 0.5-0.8 mm thick. Self-compatible; autogamous in the complex heterozygotes, outcrossing in those chromosomal homozygotes in which the stigma is elevated above the anthers. Gametic chromosome number, n=7 (7 bivalents*, ring of 14** or intermediate configurations at meiotic metaphase I). Flowering time: Octo- ber-March. Lectotype: Jacquin, Icon. Pl. Rar. 3: tab. 456. 1795. Cultivated in Vienna, seeds from Port Desire, (“Champion River"), Patagonia, Argentina, collected by Capt. Middleton and sent to Jacquin's son in 1793 by Sir Joseph Banks. Two specimens collected by Capt. Middleton are in the Forsyth Herbarium (NY), one from Port Desire, the other from Bay of San Dondo. Jacq., Coll. 107. 1796; Edwards, Bot. Reg. 2: tab. 147. 1816 Distribution (Figs. 227, 242): Low elevations from sea level to 1,000 m alti- tude, in the provinces of Mendoza, Buenos Aires, Córdoba, La Pampa, Neuquén, Río Negro, Chubut, and Santa Cruz; ascending to 2,800 m elevation only in the cordilleras of Mendoza. In southern Chile it occurs only in the immediate vicinity of the border with Argentina. 5 8 from cultivated plants: A. Se ve El Peral, ca. 6 km NW of Tupungato, Santarius 1574* (ring of 4. 5 bivalents) DU M, MO). Slopes of the Precordillera near Villavicencio, 2,500 m, Santarius 1585, 15 E 1585 also M, MO); 2,300 n E Santarius 1614, 1624* (ring of 6, ring of 4, 2 Divalent), 1626, 1628, 1632, 1634 (ring of 10, 2 bivalents), 1637 E i 10, 2 danse (D 1624 also CTES, M, MO); 1,750 m, Santarius 1650 (ring of 6, 4 biva- lents), 1651** qu 1650 also MO). Dep. Las Heras, Car es de Villavicencio, ellen id 9073* (DUSS, M). BUENOS AIRES: Dunes near Mar del 9 59 Santarius 329* (ring of 4, 5 bivalents), 331, 333, 335, 336 (DUSS; 329 also CTES, M; 329, 331, 335 also MO). NEU- QUÉN: Rocks ca. 500 m E of Piedra del Aguila, 600 m, Santarius 613 (ring of 8, 3 bivalents), 616 (ring of 8, 3 bivalents), 617*, 621*, 622, 624*, 625, 626, 627 (ring of 4, 5 bivalents), 629, 630*, 633*, 634, 635 (DUSS; 616, 617, 630 also CTES; 617, 621, 633 abu M; 616, 617, 624, 626, 630 also MO). Río Limay at Ruta 237 ca. 75 km SSW of Pie dra del Águila, San- tarius 644* (DUSS). Bank of Río Limay near Nahuel Huapí, 750 m, Santarius 879**, 882* 884 (DUSS). rio Necro: Bank of Río Limay at leaving of the Lago Nahuel Huapi, 750 m, Santarius 887, 889, 892, 893** 899, 902 (DUSS). Sandy and waste places along shore of 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 497 Lago Nahuel Huapi near the railroad station of San Carlos de Bariloche, 780 m, Santarius 653 (ring of 12, 1 bivalent), 664 (ring of 4, 5 bivalents), 666, 668, 669 (ring of 10, 2 bivalents), 670* (ring of 10, 2 bivalents), 679 (ring of 6, 4 bivalents), 680* (ring of 6, 4 bivalents; ring of 8, 3 bivalents; ring of 10, 2 bivalents), 681, 682 (ring of 12, 1 bivalent), 686*, 688 (ring of 12, 1 bivalent), 689, 690, 693, 696*,** (ring of 10, 2 bivalents), 697* (ring of 12, 1 biva- 4, 5 bivalents) 723 (ring of 8, 3 bivalents ), 724*. 726* 727, 732 742, 746 de of 10, 2 bivalents), 22 (ring of 10, 2 bivalents), 751* (DUSS. 680, 716 also CTES; 653, 680, 689, 716 also M; 680, 686, 716, 746 also MO). E slope of Cerro Otto, 1 km W of Bariloche; 850 m, Santarius 752*, 754, 755, 760 (ring of 10, 2 bivalents), 761 (ring of 6, ring of 4, 2 biva- leuts). 762, 765 (ring of 10, 2 bivalents), 766, 767* (ring of 10, 2 bivalents), 770, 771 oe. of 6, 4 bivalents), 777, 783** ( DUSS; 752 also CTES, M, MO); 950 m, . 788 * (ring of 8, 3 bivalents) (DUSS). Slopes near Ruta 258, ca. 3 km N of Rio Villegas, 67 dn Í of Bariloche, 700 m, Santarius 800* (ring of 4, 5 bivalents), 801 (ring of 6. 4 bivalents), 802, 803, 806, 810**, 812, 814, 816, 819, 820 (ring of 8, 5 (DUSS; 800 also M; 800, 801, 810 also CTES, MO). Estancia San Ramón,” W slopes at Rio Limay, E of the bridge of Ruta 237 across the river, 750 m, Santarius 854 (ring of 6, 4 bivalents; ring of 8, 3 biva- lents; ring of 12, 1 bi tient), 866**, 868**. 872, 876, 878 (ring of 12, ent (DUSS; 878 also CTES, M, MO). CHUBUT: "Sandy 3 vaste places in Villa Balneario Rada Tilly, ca. 14 km S of Comodoro Rivadavia, Santarius 937 (ring of 12, 1 bivalent), 938 (ring of 12, 1 biva- lent), 945 (ring of 12, 1 bivalent), 946, 947 (ring of 12, 1 bivalent), 948, 950 (ring of 12, bivalent), 951, 953*, 954 (ring of 12, 1 bivalent), 957 (ring of 12, 1 bivalent), 958 (DUSS; 938, 950, 953, 957 also CTES, M, MO). SANTA CRUZ: Sandy p laces S of Puerto Deseado, at mouth Ht Río "Deseado; Santarius 959* *, 961**, 963-967, 973**, 975, 979, 981**, 982**, 988 (ring of 8, 3 bivalents), 991, 994 (DUSS; 959 also CTES; 959, 982 also M; 959, 973, 981, 982 also MO). Sandy Haos in steppe with Stipa, 2 km NNE of Calafate at Lago Argentino, Santarius 1162 (ring of 6, ring of 4, 2 bivalents; ring of 8, 3 bivalents), 1163, 1166, 1171 (ring of 4, 5 bivalents), 1175, 1179, 1182 (ring of 8, 5 bivalents), 1185 (ring of 6, 4 biv: alent), 1190 nue of 6, 4 bivalents), 1193 (ring of 6, ring of 4, 2 bivalents), 1194, 1196 (ring of 4, 5 biva- lents) (DUSS; 1179 also M; 1175 also MO). Slope W of Arroyo Calafate, 350 m, Sontantus $28 CORE of 8, 3 bivalents), 1206, 1208*, 1210, 1211 (DUSS; 1208 also M; 1208, 1210 also O). Sandy places at E edge of Calafate, 200 m, Santarius 1213*, 1216 (ring of 4, 5 biva- » nts), 1217, 1218 (ring of 8, 3 bivalents), 1220, 1228* (ring of 6, "ng of 6, 1 bivalent), 1230* (ring of 4, 5 bivalents), 1233, 1239, 1241 ( DUSS; 1213, 1230 alsd M; 1213, 1241 also MO). Near the airport of Calafate, 200 m, Santarius 1245*, 1247, 1249, 1252 ( DUSS; 1245 also CTES, M, MO). Stony and sandy pla ices in the vicinity of Perito Moreno, 380 m, Santarius 1257 (DUSS, MO). Lago Buenos Aires, at Rio Los 3 W of Los Antiguos, 220 m, Santarius 1322*, 1329 3o of 12, 1 bivalent). 1331, 1332*, 1335 (ring of 10, 2 bivalents), 1336* (ring of 6, 4 bivalents; ring o of 8, 3 bivalents; ring of 10, 2 bivalents) (DUSS; 1322, 1332, 1335, also M; 1329, 1335, 1336 also CTES, MO). CHILE. MAGELLANES: Ultima Esperanza near Salto de Paine, Moore E (DUSS, M, MO). 0-4 km E of Puente Lago Amarga, W of Lago Sarmiento, S of Lago Nordenskjold, Santarius 1048**, 1049 (ring of 12, 1 bivalent), 1050 (ring of 12, $ d rem 1052**, 1054, 1055, 1058** (ring of 12, 1 bivalent), 1060, 1061 (ring of 12, 1 bivalent), 1064** 855 1048, 1050 also CTES, M; 1048, 1050, 1058 also MO). Representative specimens examined: ARGENTINA. SAN JUAN: pus 5 ( SI-4859). MN DOZA: Villavicencio, 2,000 m, Arque & Barkley in Fui (L NY); Sleumer 469 (B, LIL), Roig 5304 ( CORD); Ruiz Leal 1084 (Leal, LIL, POM); ia 9237 (BAB); Wall in 1946 (S); Mexia 04386 (GH, MO, UC); Bartlett 19403 (GH, MICH, SI, US); Senn 4443 (RSA). Los Hornillos, 2,600 m, Barkley & Paci 240 (LIL, NY). Puente del Inca, 2,800 m, Wall in 1946 (S). La Polvareda, 1850 m, Palacios & Barkley 20 Mz331 (LIL, NY ). Tony ie San Pablo, Ruiz Leal 1812 (Leal, LIL, POM). Quebrada del Arroyo Manzano, 1,650 m, Roi ig 4657 (CORD). Dep. Malalhue, Potimalal, d 500 m, Ruiz Leal 7492, (Leal, LIL). Dep. Luján, El Carmelo, 2,000 m, Cuezzo 2615 (LIL). Dep. San Rafael, Río S: dada. Ciénaguita, Rossi 294 (LIL). Dep. Tupungato, Canal 5 tir near Toma, Ruiz Lea l si Ces l, LIL, LP). Near San Pedro, Gillies in 1824 (K). Paso Cruz, Kuntze 59 (CORI „F). . San Carlos, Quebrada Alvarado, Covas 3492 (SI). Laguna Carrilauquen, Kurtz 755 jean. MO). Atuel Valley near El Sosneado, 1,600 m, Bócher & Hjerting 881 (C). NguqQquÉN: Pulmaré, Comber 388 (K). Between Catanlil and Junin de los Andes at Rio Aluminé, Bécher et al. 1643 (C). Parque Nahuel Huapi, De Barba 2106 (LIL, RSA); Jacobsen 15 (POM). Manza near Zapala, Ancibor 90245 (BAB). 498 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Cohün-Có, Comber 869 (K). Lago Huechulafquen, O'Donell 2320 (LIL, NY, S, SI). Junc- tion of Río Limay and Río Traful, Cardini 46 (LIL). Cerro Lotena, 900 m, Ammann 82 (F, ío Barrancas at Ruta 40 near Pehuenches, Ancibor 90134 (BAB). San Martín de los Andes, Dawson 1297 (BAA). Lago Lolog near San Martín de los Andes, Scolnik 230 (RSA). RIO NEGRO: Parque Nahuel Huapi, De Barba 750 ( BM, LIL), 1462 ( LIL, RSA); Cardini 228 (US); Giovanelli 13600 (BAB); Bernicken in 1896 (LP), Arnow 3761 (MO); Fabris 1124 (BR); Buchtien 1356 ( AMD, BREM pro parte, GH pro parte, L, LE, LIL, LY, M, S, SI, US, W), 17 (POM). Banks of Río Negro near General Roca, Krapovickas t Cristóbal 22414 (MO). Between Laguna de las Banduras and Fortin Fé, N.N. in 1879 (CORD). El Condor near Viedma, Cabrera et al. 19561 (LP, P). Carmen de Patagones, Haumann in 1912 (BA). Dep. San Antonio, Piccinini 1291, 1415, 1485, 1833 ( BAB); Hicken 37 (SI). Estuary of =y Negro, N.N. 86 (LE). CHupur: Colonia Sarmiento, O'Donell 3485 (LIL); Cabrera 45 (LP Puerto Madryn, Dusén 5323 (SI); O'Donell 3255 (LIL); Soriano 2713 (BAB). Colonia San Martin, Gerling 31 (POM). Gobernador Costa, Birabén 580 (LP). Camarones, sea shore, Aurelius 25 (S). Peninsula Valdez, Rovereto 31-1537 (POM). Escalante, O 'Donell 3551 (LIL); Kreibohm 110 (LP). Bahía Camarones (Port Sta. Elena), Anderson 24 in 1826 (BM). Valley of Laguna Blanca, Koslowsky 201 (BM, K, SI, Z). Esquel, Castellanos in 1945 (F, LIL); Eyerdam et al. 24575 (G, K, UC); Kühnemann 647 (RSA). Corcovado, Illin 6872, 6876 (BAB), 94 (BR, CORD, HBG, SI); Soriano 3028 (CTES). Lago Futalaufquen, Correa 4153 (BAB, UC); Constance et al. in 1967 (BAA, MO); Hicken 4, 14 (SI), El Maitén, Meyer 9705 (LIL). sanra cruz: Vicinity of Lago Argentino, Dusén 5787 (S, SI); Sleumer 1238 (LIL, US); Boelcke 12509 (BAA, BAB); Eyerdam et al. 24274 (G, GH, K, MO, SI, UC); Scolnik 368 ( a iri 99a (GH, US), 5322 (NY); Koslowsky 72 (CORD). Tehuel- ches, Donat 54 (B GH, HBG, K, LIL, MO, NY, S, SI, UC, Z). Lago San Martin, Rohmeder in 1945 P III.). San Julián, Blake 56A (LIL). Río Coyle, Dauber 84 (POM ). Rio Gallegos, Brown 62 (NY). Puerto Deseado, Ancibor d» Vizinis 4405 (BAA, MO); ~ 12145 (BAA, BAB); Eyerdam et al. 23869 (G; GH, K, MO, SI, UC), O’Donell 3630 (LIL); d 49 in 1826 (POM). cónpoBa: Achiras, Trelles (SI-4853). LA PAMPA: Cerro Lihuel Calel, 400 m, Burkart 20555 (P, SD; Schwabe i Fabris 2028 (LP); Krapovickas 3620 (BAB), P ceci & Cristóbal 22384 (MO); Troncoso in 1959 (SI); Boelcke d» Nicora 8120 (BAA, O). General Acha, Monticelli 10 (SI); Troncoso in 1959 (SI). BUENOS Amts: Balcarce, D 3828 (POM). San Clemente near General Lavalle, Cabrera 4921 (GH). Sierra La Tinta, Spegazzini 40 (BAB). Quilmes, Hicken in 1904 (SI). Mirimar, Cabrera 5559 (LP). Mar del Sur, use 17876 (SI); Nicora 17876 (SI, US). Pehuén-Có, Correa 2299 (BAB); Cabrera 14911 (LP, M); Erettowi 2732 (MO); Boelcke 11964 (SI). Bahía Blanca, Ameghino 31-1628 (POM); Job 1599 (NY). Bahia San Blas, Fabris & Schwabe 5016 (CTES, M). Sierra Curamalal, Spegazzini 18 (BAB); Hohnberg in 1884 (CORD); Burkart 4796 (BAA, CTES, MO); Cabrera 5496 (L.P); Parodi 10355 (BAA, POM). Monte Hermoso, Alboff in 1916 (LP); Fabris & Schwabe 4817 (M); Cabrera et al. 17055 (LP); De Barba 633 (LIL); Eskuche & Klein in 1968 (CTES). Quequén, Castellanos 16092 (LIL); Dawson 662 (LP, NY); Rodri- guez 828 (GH, LIL, NY). Necochea, Nicora in 1961 (BAA), 7047 (MO); Fabris & Schwabe 4759 (CTES, M, RSA); Rodriguez 848 (GH, NY); Eyerdam et al. 23709 (G, GH, K, UC). Sierra de la Ventana, Alboff 124, 343 (CORD); Cano & Cdmara 227 (BAA, BAB); Dawson 131 (LP), Krapovickas 2958 (LIL, MO, RSA, SI); Pastore 1205 (F, LIL, SI); Abbiatti 4253 (NY). Olavarria, Cabrera 20929 (P). Sierras del Azul, Osten 152 (BREM). CHILE. AISEN: Coihaique, Rentzell 6129 pro parte (G, SI). Mina Silva near Lago Buenos Aires, Heim in 1939 (Z). MAGELLANEs: Puert i. 1 106 (SI). wq from plants cultivated in garden . Kew, seeds from Port Desire, 5 in 1791 (BM). Botanical Garden Paris, in 1814, Bt Cambessedes (MPU). Botanical Garden Munich, Germany, in 1814, Herb. Mar- tius (BR). Botanical Garden Arlary, in 1827 (GH). Botanical Garden Munich, in 1832 (M as O. undulata Horti). Munich, in 1833, (BR; as O. undulata). Warsaw, Poland, in 1834 (LE; as O. undulata). Botanical Garden Paris, in 1842 (BR). Leningrad, USSR, in 1847 (LE; as O. dubia F.M.). Leningrad, in 1848 (LE; as O. cognata). Botanical Garden of Vienna, seeds from Berlin, in 1856 (W; as O. villosa). Cultivated at Nymans, Sussex, England, 1929, Comber 388 (K). Cultivated in Botanical Garden, Leningrad, seeds from D. Nolte, 1847 (LE; as O. dubia F.M.). Cultivated in Botanical Garden: Leningrad, 1847 (LE; as O. cognata). Oenothera odorata reported in the literature from outside of South America: GERMANY: Hegi (1925; pt. 2: 864). DIETRICH—SOUTH AMERICAN OENOTHERA Ficures 140-143. Taxa of Oenothera sect. Oenothera subsect. Munzia.—140. O. picensis 1 ied (Argentina, Mendoza, 5 1554).— 141. O. picensis subsp. cordobensis (Argentina, Jujuy, Santarius 1 . picensis subsp. bonariensis (Argentina, Buenos Aires, pis 370).—143. O. M tectis (Uruguay, Montevideo, Santarius 1). 500 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Oenothera odorata is characterized by its large flowers with a relatively short floral tube; long, narrow fruits; and bracts broadly rounded at the base. Aside from O. magellanica, it is the only frequent member of the subsection in southernmost South America. This species is extraordinarily variable, but it is impossible to arrange its variability into an inclusive and useful taxonomic sys- tem; consequently, no infraspecific taxa are recognized. Especially pronounced is the variability in habit, hue of the stems and leaves, leaf shape, and pubescence. Plants with plane leaves grow intermixed with others with somewhat undu- late leaves in the same populations, in which intermediates also occur. Reddish or greenish coloration of the above-ground parts segregates in a Mendelian fashion, as it does in other species of the genus. Plants with strigillose pubescence are completely absent in the northeast part of the range, in the province of Buenos Aires, but are widespread else- where. They predominate in the south, whereas populations consisting exclu- sively of plants with villous pubescence are more frequent in the north. Popu- lations of this species in the province of Buenos Aires consist mainly of plants with an almost woolly pubescence, and are distinctive in this respect. Such populations have not been accorded taxonomic recognition, however, because they include, in addition to those with woolly pubescence, others which are identical to the less densely pubescent forms that are frequent in the provinces of La Pampa, Neuquén, and Rio Negro. The hairs that make up the so-called strigillose pubescence of O. odorata are not typical of those normally included in this category of pubescence in the species of series Renneria. In this series strigillose hairs are ca. 0.2 mm long and appressed closely to the stem, whereas in O. odorata they are about twice as long and generally erect, with only the tip curved toward the stem. The pubescence in O. odorata could therefore with justification be regarded as inter- mediate between strigillose and villous in character. Oenothera odorata is extremely variable in its chromosomal configurations. Plants with 7 bivalents and all configurations up to and including complex heterozygotes with a ring of 14 occur. In the vicinity of Comodoro Rivadavia, Chubut Prov., the essentially stable configuration ring of 12 + 1 bivalent seems to have become predominant. This situation has been discussed further on p. 437. 16. Oenothera ravenii Dietrich, sp. nov.—Fics. 6, 41-43, 47-49, 124-127, 176, Herba annua vel biennis, erecta, rosulata, simplex vel uen principalis ramosus et ramis plerumque late arcuate vel rariore oblique e rosula ascendentibus, 5— m alta. Plantae dense villosae, praecipue inferiore, sparseque 5 pubescentes 2 ce ad sparse strigulosi praecipue inferiore. Folia rosulae cultrata vel anguste oblanceolata, UE breviter acuta, € acuta vel trun cata, sess silia, 5-20 c cm lon nga, Pic cm Bec folia caulina cultrata ad c 502) em lata; b oe ovata vel masaq acts. basi truncata vel subcordata, ses- silia, plerumque quam capsulam subtena multo breviora, raro ad eam subaequalia, 1.5-3 cm longa a, 0.5-1.5 cm lata; folia subintegria vel irregulariter obtuseque serrata, saepe marginibus praecipue in bracteis rubrescentibus. Inflorescentia simplex vel ramosa. Tub s floralis (2— 5.5(-6.5) cm longus. Gemmae ambito ese vel lanceolatae, 1-3.5 cm longae, 5-11 mn DIETRICH—SOUTH AMERICAN OENOTHERA ae is s O 1 Ficut 144-147. longiflora n Arge ntina, ( Uruguay [Taxa of Oenothera sect. Buenos Aires, Florida, Santarius 211 Aires, Santarius 468) 23423). Oenothera subsect. Munz Santarius 27 146. 147. a—144. O. seudo- aue ani subsp. parodiina 0 parodiana (Argentina, O. parodiana mbs. brasiliensis ( Argentina, Entre Rios, Burkari 502 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 crassae, plerumque junctura sepalorum tubo florali rubro-fasciatae, flavae vel flavido-virescen- tae, saepe rubrae. Sepala saepe dense vel sparse rubro-maculata; apices sepalorum erecti vel divergentes, 1-3 mm longi. Petala latissime obovata, lutea, saepe basi rubro-maculata, 1.2— (-5.5) cm longa. Stylus longus, stigmate sub anthesi supra antheras elevato, vel brevis, stig- mate sub anthesi antheris circumdato. Ovarium 1.5-2.5 cm longum. Capsula 2.5-3.5 cm longa, 3-4 mm crassa, plerumque ab caulo directo subperpendicularo curvatum. Semina am- bito elliptica vel rotundata, fusca, 1-1.5 mm longa, 0.5-0.7 mm crassa. Numerus gameticus — chromosomaticus, n = 7; planta chromosomatice homozygotica, heterozygotica complexa, vel intermedia Erect annual or biennial herb, forming a rosette, unbranched or with a branched main stem and side branches which are usually widely arcuate but sometimes obliquely ascending from the rosette, 5-10 dm tall. Plants + densely villous, especially below, and sparsely glandular-pubescent or moderately to sparsely strigillose, especially below. Rosette leaves cultrate to narrowly oblan- ceolate, mostly short acute, acute to truncate at the base, sessile, 8-20 cm long, 1.5-3 cm wide; cauline leaves cultrate to narrowly oblong or lanceolate, acute, truncate to subcordate at the base, sessile, 3-15 cm long, 0.8-1.5(-2) cm wide; bracts narrowly ovate to ovate, acute, truncate to subcordate at the base, sessile, mostly much shorter than the capsule they subtend, rarely + the same length, 1.5-3 cm long, 0.5-1.5 cm wide; leaves plane or undulate at the margins, subentire or irregularly serrate with blunt teeth, often reddish along the mar- gins, especially the bracts. Inflorescence branched or unbranched. Floral tube (2-)3-5.5(-6.5) em long. Buds oblong to lanceolate in outline, 1-3.5 cm long, 5-11 mm thick, usually reddish at the junction of the sepals with the floral tube, yellow or yellowish green, often flushed with red. Sepals often densely or sparsely flecked with red; apices of the sepals erect or divergent, 1-3 mm long. Petals very broadly obovate, yellow, often with a red spot at the base, 1.2-5 (-5.5) cm long. Anthers 6-13 mm long. Filaments 8-27 mm long. Style long, the stigma held above the anthers at anthesis, or short, the anthers shedding pollen directly on the stigma at anthesis, 3-9 cm long. Stigma lobes 4-9 mm long. Ovary 1.5-2.5 cm long. Capsule 2.5-3.5 cm long, 3-4 mm thick, mostly curved directly outward from the stem. Seeds elliptic to rotund in outline, brown, 1-1.5 mm long, 0.5-0.7 mm thick. Self-compatible; outcrossing in those plants, all chromosomal homozygotes, in which the stigma is held above the anthers at anthesis, and self-pollinating in the rest. Gametic chromosome num- 7 bivalents, ring of 14 or intermediate configurations at meiotic meta- phase I). Type: Grown from seed and cultivated in the Botanical Garden of Diissel- dorf, Germany, 15 Aug. 1972. Source: Brazil, State of Rio Grande do Sul, Pelotas, 1966, E. J. Hackbart (MO-2155712, holotype; CTES, DUSS, M, isotypes). Distribution (Figs. 225-226, 228, 243): In Brazil from Rio Grande do Sul north to the state of Sao Paulo; in Uruguay in the provinces of Salto, Cerro Largo, Rocha, Lavalleja, and Montevideo; in Argentina in the provinces of Misiones, Corrientes, Entre Rios, Santa Fé, Buenos Aires, and Córdoba; in cen- tral Paraguay; and in Chile from Valdivia to Valparaiso. This new species is dedicated to Peter H. Raven (1936-). Among its char- acteristics are the short, red-margined bracts and the sessile, usually oblong 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 503 rosette and cauline leaves. Within this species subsp. ravenii is chromosomally homozygous, whereas subsp. argentinae and subsp. chilensis are entirely chro- mosomally heterozygous, but deriving both of their complexes from O. ravenii. Hybrids of chromosomal heterozygotes with homozygotes within this species yield only two closely similar phenotypes, both easily assignable to O. ravenii, in the F, generation. Similarly, hybrids with other homozygous species show that all elements included here within O. ravenii agree closely genetically de- spite their chromosomal differences. KEY TO THE SUBSPECIES l. Petals 2.5-5 cm long; stigma usually elevated above the anthers at anthesis; buds 2- 3.5 cm long ja. subsp. ravenii l’. Petals 1.2-3 cm long; stigma surrounded by the ec E anthesis; buds 1-2 cm lon 2. diee lanceolate in outline, 1-1.7 cm long; petals 1.2-2 cm long; seeds 1-1.3 mm 6b. subsp. idees E oblong in outline, 1.5-2 cm long; petals 2— 2.5 cm n long; seeds 1.3-1.5 1 lona Lon 2 ĩͤé-/ A RS MU l6c. subsp. cr, 16a. Oenothera ravenii subsp. ravenii—F ics. 6, 41-43, 124-125, 176, 212. O. mM L. var. poraguayensts Chod., Bull. Herb. Boissier 7(9, app. 1): 71. 1899. LEC- PE: Paraguay, margins of fo rests near Cordillera de Altos, July (1885-1895), E. Hassler 338 (G, holotype; G, P, isotypes); Hassler, Bull. Soc. Bot. Genéve, sér. 2, 5: 27 1913 O. mollissima sensu Chodat, Bull. Herb. Boissier 7(9, app. 1): 5 1899. O. longiflora sensu Munz, Physis 11: 285. 1933, pro parte; Amer. J. Bot 663. 1935, parte; Comun. Bot. Mus. Hist. Nat. Montevideo 1(10): 35. 1943. pro sei Fl. Brasílica 9(41): 1947. O. parodiana sensu Munz, Amer. J. Bot. 22: 662. 1935, pro parte. Rosette leaves cultrate to narrowly oblong or oblanceolate. Floral tube 3- 9.0(-6.5) cm long. Buds lanceolate in outline, 2-3.5 em long, 7-11 mm thick. Sepals reddish around the edges and often flecked with red on the surface; apices of the sepals 2-3 mm long. Petals 2.5-5 cm long. Anthers 8-13 mm long. Filaments 18-27 mm long. Style long, rarely short, the stigma usually elevated above the anthers at anthesis, 5.5-9 cm long. Stigma lobes 5-9 mm long. Ovary 1.5-2.5 cm long. Seeds 1.3-1.5 mm long, broadly elliptic in outline. Self-com- patible but mostly outcrossing or self-pollinating and complex heterozygote. Gametic chromosome number, n= 7 (7 bivalents*, ring of 14** or small rings at meiotic metaphase I). Flowering time: October—June. Distribution (Figs. 225, 243): Occurs in Brazil from Rio Grande do Sul north to Minas Gerais; in Uruguay in the provinces of Salto, Cerro Largo, Rocha, Lavalleja, and Montevideo; in central Paraguay; and in Argentina in the prov- inces of Misiones, Corrientes, Entre Rios, and Santa Fé. Specimens examined from cultivated pla RAZIL. RIO GRANDE DO SUL: Pelotas, Hackbart 1966* (ring of 6, 4 bivalents; ring of 8, ring of 4, 1 5 (CTES, DU SS, M, MO). SANTA CATARINA: Coast near Ararangua, Schultz | in 1970 (ring of 4, 5 bivalents ) (DUSS). NTINA. 7 8 At Ruta 12 near San Ignacio, Conrad & Dietrich 26, 27 (ring of 6, 4 1 24 (2 rings of 6, 1 bivalent) (DUSS). CORRIENTES: Est. Garruchos near Santao Tomé, Krapovickas 21464** (DUSS) 504 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ficures 148-151. diana subsp. strigulosa (Argentina, Buenos Aires, Spegazzini in — 540 ).— Peru, Arequipa, Munz . - T Oopel inet metase la Breve rot Basis. aman MOA Airs, anan, for AMerilla. ba [Vor. 64 O. verrucosa Taxa of Oenothera sect. he dila subsect. Munzia.—148. O. paro- 1938 149 O. 5 (Chile, Atacama, Jiles 2160). — 151. O. coquimbensis (Chile, Atacama, Johnston 4990). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 505 GUAY. CORDILLERA: Roadside near Altos, ca. 10 km NE of San Bernardino, Conrad e Dietrich 30**, 36** (DUSS). Ca. 4 km SSE of San Bernardino, Conrad H Dietrich 39** USS). CENTRAL: 3 km S of K un Conrad & Dietrich 50**, 53** (DUSS). oo per s? examined: L S IS: High fields in the Serra do Pieü, 1,800 m, Schwacke 5248 (BR). SÃO PAULO: st. "Hilaire ges in PARANA: Eng. Bley near Lapa, Hatschbach 1067 (S, US). Between Jaguariaiva and Senges, Hatschbach 9064 (HB, L). Roca Nova near Piraquara, Hatschbach 1898 Da Pe mta Grossa, Schwacke in 1874 (R); Hoehne 23248 EE Curi- tiba, Galvão in 1884 (R); Stellfeld 1112 (SP). 50 km W of Guarapuava, 1,000 m, Reitz & Klein 17713 (US). Agua Sta. Clara near Guarapuava, Pereira 7967 (HB). Porto E Gurgel in 1929 (BR); Gimpel 16163 (BR). Vila Velha, Pereira & Pabst 7547 (M); Dusén in 1904 (S); N.N. in 1904 (R); Hertel 52325 (SP); Pereira 8272 (BR, HB). Santa Catarina: Bom Retiro, 1,650 m, Smith & Klein 10435 (HBR, RSA, US). Ponte Alta near Curitibanos, 800-900 m, Smith & Klein 8246 (HBR, NY, R, RSA, US). Agua Doce near Rio Chapeco, 12 km S of Haran Smith & Klein 13550 (HBR, RSA). Fazenda de Laranja, S. Joaquim near Bom Jardim, 1,400 m, Reitz & Klein 7735, 7977 (HBR). Palhoca near Maciambu, Reitz d Klein 980 (HBR). Garopaba, Klein & Bre solin 8855 (HBR). Palmas near Joaçaba, 52 km W of Caçador, 1,000-1,300 m, Smith & Reitz 9153 (US). Cangicas near Araranguá, 50 m, Reitz C254 (BR, HBR). Sombrio near Araranguá, 10 m, Reitz C1308 (GH, HBR). Campos Novos, 1,000 m, Klein id (HBR, RSA). RIO GRANDE DO SUL: Montenegro, Henz 32563 (LIL Uagusiami. Palacios d» Cuezzo 210 (LIL). Between Cacapava do Sul and Morro Perao, Palacios & Psa 1449 (LIL). Between Alegrete and Capivari, Palacios & Cuezzo 1919 (LIL). Itapoan, Rambo 44448 (LIL). Near Vire ae Rambo 42543 (LIL). Cerro de San — near bie Maria, Vidal 1508 m o Pardo, Vidal 01570 (R). Between Capão and Osório, Nelson in 1970 (BR). Caaró near Sió Luiz, Rambo 53349 (B). Lagoa Vermelha, ornant 9078 (MVFA). Est. do Jaran near Quarai, ee 26320 (LIL). Passo do Socorro near Vacaria, Rambo 51383 (HBR). Pelotas, Beetle 2236 (US); Costa Sacco 690 (F, HB, R). Piratiny near Pelotas, Malme 159 (S). S. Leop olde, pe 293 (LIL). Porto Alegre, Molfino in 1898 (POM); S. Barbara in 1835 (BR); Rambo 27040 (LIL). Gloria near Porto Alegre, Bornmüller 47 ( GH); Rambo 6461 URUGUAY. CERRO LARGO: Rio Branco, Herter 2104 (F, MO, NY, US, Z). Bañado de los Burros, Flossdorf 1 (POM, A Arroyo Zapallar, Praderi 742 (LIL). SALTO; Thermal Springs of Arapey, Rosengurtt 10569 (MVFA). Paso del Arroyo Las Cañas, Rosengurtt BI058 (POM). ROCHA: Castillos, Herter 987938 (POM). E coast of Uruguay, St. -Hilaire 2213 (P). Lava- LLEJA: Road to Puma near Minas, Osten 4490 (G). C. Verdün near Minas, Berro 6246 (MVFA). MONTEVIDEO: Punta Gorda, Osten 22088 (GH). ARGENTINA. MISIONES; Candelaria, Bertoni 2463 (LIL); Descole 3270 (LIL); Sesmero 83 (LIL). Loreto near Candelaria, 290 m, Montes 58B (US). At Ruta 14 near Arroyo Garupá Norte, Krapovickas et al. (CTES). GCarupá near Candelaria, Bertoni 4734 (LIL). Pindapoy near Candelaria, Bertoni 3862 (LIL). P xj Ana, Bertoni in 1944 1 * Sesmero 168 (LIL, NY); Rodriguez 671 (A LIL, POM, SI, UC); Schw wi 632 (LIL, NY, UC); Montes 1516 (RSA); Rodriguez 158 (LIL). San José, Bote 241 (LIL). Posadas, ; Galindo 3739 (SI); Bertoni 1514 (LIL). A me near 5 130 m, Zu 5724 (LIL). Posadas, Ekman 2099 (G, LD, S, US). Cerro C Crovetto 9492 (BAB). Puerto Leoni near Cainguas, Schwarz 1575 (LIL). Apostoles, Ibarrola 1051 (BR, LIL, NY). Santa Irene near San Javier, Bertoni 526 (LIL). Corpus near Santa Ana, Bertoni 1866 (LIL). Sto. Tomás, 160 m, Bertoni 4696 (LIL). Santa Inés, Meyer 11439 (LIL). Concepción, Schwarz 3566 (LIL, RSA). Ar- royo Jabebiri, Meyer 11512 (LIL). Villa Samis, N.N. in 1944 (LIL). El Dorado near Iguazú, Schwartz 2042 (LIL). Puerto Wanda near Iguazú, Montes 9591 (LIL). Puerto Istuela, Montes 10117 (LP). Arroyo Apepú near San Ignacio, Schwarz 2882 (LIL). Menóchio near San Ignacio, Schwarz 1255 (LIL). San Ignacio, Medina 212 (LIL, RSA). CORRIENTES: Es- ancia “Santa Teresa,” Pedersen 93 pro parte (S, US). Est. “Garruchos” near Santo Tomé, 10 ge 9229 (C); 1 et al. 21464 (MO). At Ruta 14 2 Gob. Mess aeo ickas et al. 16692, 16753 (CTES). Est. San Francisco, 23 km NW of Gob. Vira sebo et al. 17217 (CTES). Paso Pucü near Concepción PUE 7482 (C). "Balbiese, Castellanos 34445 (RSA). S. Roqueito near Mercedes, Irigoyen 71 (CTES). San Carlos near Ituzaingó, Krapovickas et al. 17995 (CTES). Yahápe at Ruta 12 near Berón de Astrada, Kra- povickas et al. 16537 (CTES). ENTRE Rios: Concepción del Uruguay, Lorentz in 1876 (BM). SANTA FÉ: Road to Requoncista, Job 974 (NY PARAGUAY. Cordillera de Altos, Fiebrig 434 (F, G, GH, HBG, K, L, LY, M); Hassler 338 (G, P), 744 (G, K, P). San Bernardino, Hassler 3343 (BM, G, GH, K, LY, NY, P, W). Near 506 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Igatimi, Hassler 5567 (BM, C, GH, K, LIL, LY, MO, vua P, S, UC, W); Tóppen in 1883 (H Vicin i^ = the river Y-aca, Hassler 6879 (BM G, GH, K, LIL, LY, MICH, MO, MPU, NY, P, S, UC, W). Vicinity of the lake Ypacaray, Hassler 12486 (BM, C, F, G, GH, K, L, LIL, LY, MICH, MO, NY, S, UC, US, Z). Villa Elisa, Pedersen 5125 (C). Caáguazuü, Balansa 2220 (G, P, RSA); Hassler 8996 (BM, G, K, NY, W); Tóppen in 1883 (HBG). Tapyta, Jórgensen 4754 (F, MO, NY, POM, S, SI, US). Sapucaí, Gótzsche in 1891 (CORD). Pirareta near Paraguari, Sparre "m Vervoorst 173 (LIL). Chololo near Paraguari, Sparre & Vervoorst 613 (LIL). Encarnación, 110 m, Bertoni 4524 (LIL); Schrottky 116 (LIL). Early specimens from plants cultivated in Botanical Gardens: Rovig in 1818 (CORD; as O. longiflora). From the Garden of Mr. Ohm at Berlin, Bauer in 1838 (CORD; as O. sellowiana). From Botanical Garden at Berlin, Bauer in 1842 (CORD; as O. sellowiana). Oenothera odorata and O. featherstonei are the only entities that regularly exceed O. ravenii subsp. ravenii in flower size. In this subspecies, plants with red-flecked and entirely green sepals grow intermixed in the same populations, and the characteristic seems to segregate in a Mendelian fashion. Plants from Brazil, Uruguay, and the provinces of Corrientes and Entre Ríos in Argentina are moderately pubescent, whereas those from the Province of Misiones in Argentina are very densely pubescent, especially in their lower portions. On the map of distribution of the ravenii-complex (Fig. 243), it can be seen that this chromosomal complex exists in homozygous form only in the north- eastern portion of the range. Evidently, this complex combines readily with others as discussed under species no. 23, 27, 28, 36, 37 and 41. As the ancestors of O. ravenii migrated east from a probable area of origin in the vicinity of Tucumán, they seem to have hybridized readily with others, and the homozygotes persist today only at the very margins of the range. Just as in O. odorata, the existence of an array of chromosomally differenti- ated homozygotes may be inferred within this entity, the hybridization of which has given rise to the complex heterozygotes. In Düsseldorf two chromo- somally homozygous plants (72-1227a and 72-1228) from the same locality near Pelotas were crossed and gave rise in the F, generation to a plant (73-576) with a ring of 14 chromosomes, showing that such potentiality existed also within the original population. 16b. Oenothera ravenii subsp. argentinae Dietrich, subsp. nov.—Fic. 126. O. parodiana sensu Munz, Physis 11: 283. 1933, pro parte; Amer. J. Bot. 22: 662. 1935, pro parte. O. stricta sensu Munz, Physis 11: 285. 1933, pro parte. Folia ut in subsp. ravenii. Tubus floralis 1.8-3.5 cm longus. Gemmae ambito lanceo- latae, 1-1.7 cm longae, 5-7 mm crassae. Sepala iid -marginata, saepe etiam rubro-maculata; apices sepalorum 1-2 mm longi. Petala 1.2-2 c nga. Stylus us stigmate sub anthesi antheris circumdato. Ovarium 1.3-1.5 cm 1 Semina 1-1.3 mm longa, 0.5-0.7 mm crassa, ambito rotundata. Numerus gameticus chromosomaticus, n = 7, poe chromosomatice heterozygotica complexa Leaves as in dies ravenii. Floral tube 1.8-3.5 cm long. Buds lanceolate in outline, 1-1.7 cm long, 5-7 mm thick. Sepals with red margins, often also flecked with red; apices of the sepals 1-2 mm long. Petals 1.2-2 cm long. An- thers 6-8 mm long. Filaments 12-15 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 3-4.5 cm long. Stigma lobes 4-5 mm 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 507 | " ö fs 5641606 beget IGURES 152-153. Taxa of Oenothera sect. Oenothera subsect. Munzia.—152. O. arequi- pensis (Peru, Arequipa, Mexia 04167).—153. O. featherstonei (Peru, Lima, Weberbauer 5217). long. Ovary 1.3-1.5 cm long. Seeds 1-1.3 mm long, 0.5-0.7 mm thick, rotund in outline. Self-pollinating complex heterozygote. Gametic chromosome num- ber, n = 7 (ring of 14* or ring of 12 and 1 bivalent** at meiotic metaphase I). Flowering time: October-April. Type: Grown from seeds and cultivated in the Botanical Garden of Diissel- dorf, Germany, 14 Aug. 1972. Source: Argentina, Prov. Cordoba, Copina near Córdoba, Dec. 1961, G. Gópel (MO-2155709, holotype; CTES, DUSS, M, iso- types ). Distribution (Fig. 228): In Brazil, so far known only from the state of Rio Grande; the southern departments of Canelones, Maldonado, and Florida in Uru- guay; and the provinces of Entre Rios, Buenos Aires, and Cordoba in Argentina. Specimens examined from cultivated plants: AZIL. RIO GRANDE DO SUL: Pelotas, Hackbart in 1966* (DUSS, M); Hackbart in S ARGENTINA. ENTRE Rios: Concepción del Uruguay, Burkart in 1961* (DUSS). CÓRDOBA: Copina near Córdoba, Gópel in 1961* (DUSS, M, MO). BUENOS AIRES: Mar de la Plata, San- tarius 342* (CTES, DUSS, JRUGUAY. FLORIDA: Florida, Hecht 1964-31* (DUSS). Mansavillagra, Santarius 231* (CTES, DUSS, !) ; oe specimens 5 BRAZIL. RIO GRANDE DO SUL: ir Caçapava, P i Sào pa: Rosengurtt 8736 (MVFA). URUGUAY. MONTEVIDEO: 5 p illaga 721 ERU oniDA; Casuja, Borsani 5564 (MVFA). CANELONES: Arroyo Sarandi, Izaguirre i (M VFA). — 508 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ENTINA. ENTRE RÍOS: Concepción del Uruguay, Lorentz in 1876 (GH). BUENOS AIRES: Vicinity of Junin, Saint-Ives 173 (G). Sierra de la Ventana, N.N. 12789 (LP). Est. San Juan, 30 km S of Buenos Aires, Eyerdam 23037a (G, GH, K, MO, UC). 20 km N of Mar del Plata, road to Balcarce, Eyerdam et al. 23644, 23669 (GH, K, UC). "La Brava" near Balcarce, Lourteig 162 (LI L); Capurro in 1941 (LIL). Cerro El Sombrerito near Tandil, 550 m, Huidrobo 1770 ewe NY). El Dia, King 263 (BM). Delta del Parana, Rio Cabo, Burkart 5119 (BAA). DOBA: Sierra Grande, Copina, Hunziker 11439 (CORD, MO). Sierra Achala, ay 335 (P). Oenothera ravenii subsp. argentinae is a complex heterozygote which can be distinguished from the chromosomally homozygous subsp. ravenii only by its smaller flowers. The plants of subsp. argentinae seem more vigorous in growth than those of subsp. ravenii and may eventually replace them. Populations of subsp. argentinae that occur within the area of subsp. ravenii have, as a rule, larger flowers than those which occur beyond this area, in the Province of Bue- nos Aires. Populations of subsp. argentinae probably represent an early phase in the stabilization of their complex heterozygosity. 16c. Oenothera ravenii subsp. chilensis Dietrich, subsp. nov.—F ics. 47-49, 127. Folia rosulae anguste VASE a vel oblanceolata. Tubus floralis 2-3.5 cm longus. ;emmae ambito oblongae, 1.5-2 cm longae, 5-6 mm crassae, E iy a sepalorum tubo florali rubro- fasciatae. Sep ala 3 apices sepalorum 1.5-2 mm longi. Petala 2-2.5 cm longa. tor brevis, stigmate sub m antheris circumdato. Ovarium 1.2-2 cm longum. Semin: 1.3-1.5 mm | longa, € 0.5-0.6 mm crassa, ambito late elliptica. Numerus gameticus chromosomati- cus, n = 7; planta 5 heterozygotica complexa. Rosette leaves narrowly oblanceolate to oblanceolate. Floral tube 2-3.5 cm long. Buds oblong in outline, 1.5-2 cm long, 5-6 mm thick, red at the junction of the sepals with the floral tube. Sepals not red-flecked; apices of the sepals 1.5-2 mm long. Petals 2-2.5 cm long. Anthers 6-8 mm long. Filaments 8-13 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 3-5 cm long. Stigma lobes 4-7 mm long. Ovary 1.2-2 cm long. Seeds 1.3-1.5 mm long, 0.5-0.6 mm thick, broadly elliptic in outline. Self-pollinating complex heterozygote. Gametic chromosome number, n — 7 (ring of 14* at meiotic metaphase I). Flowering time: October-February. Type: Grown from seeds and cultivated in the Botanical Garden of Düssel- dorf, Germany, 24 July 1973. Source: Chile, Prov. Cautín, Molco at Lago Villa- rica, 1960, W. Stubbe (MO-2155707, holotype; CTES, DUSS, M, MO, isotypes ). Distribution (Fig. 226): In Chile from the provinces of Valdivia to Val- paraiso. 5 examine d from cultivated plants: HILE, CAUTIN: Fundo Doyin near Lonco ché, Stubbe in 1960* (CTES, DUSS, M, MO). Molco at bu. Villarica, Koch in ti (DUSS); Stubbe in 1960* (DUSS, M, MO). Additional specimens examin HILE. VALPARAISO: Valarin. Buchtien in 1895 (US). coNcgEpcióN: At km 15 be- tween Concepción and Coronel, Cea & Ugarte in 1967 (CONC-35028). At km 42 between Concepción and Bulnes, Villarroel & Weldt 119 (CO de^ iine Concepción, Macrae in 1825 . LLECO: Fundo a Elias near Victoria, 300 m, Sparre 3315 (S). VALDIVIA: Panguipulli, 230 m, 1 562 (M). Quinchiles, * 65a (LP). Oenothera ravenii subsp. chilensis can be distinguished from both other sub- species by its medium-sized flowers, buds oblong in outline, and consistently 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 509 oblanceolate leaves. It apparently arose from Chilean populations of the com- plex heterozygote O. stricta subsp. stricta by the segregation of two genomes originally derived from subsp. ravenii and the reconstitution of a chromosomally homozygous series of populations west of the Andes. In cultivated plants the influence of O. odorata on O. ravenii subsp. chilensis can be seen, especially in the form of the rosette leaves. This influence has come about because of cross- ing-over between the ravenii-genome and the odorata-genome within the com- plex heterozygote O. stricta subsp. stricta. The genomes of O. ravenii that seg- regated from O. stricta subsp. stricta apparently were modified in this way, and the resulting populations, although primarily O. ravenii, show the influence of O. odorata. In a similar manner, O. affinis seems to have modified Paraguayan populations of O. ravenii subsp. ravenii. 17. Oenothera 5 L., Mant. Pl. 227. 1771.—Fics. 50-52, 55-57, 128- 129, 177, 213-214 Erect annual or biennial herb, forming a rosette, unbranched or with a branched main stem and widely arcuately ascending side branches arising from the rosette, 4-8 dm tall. Entire plant densely long-villous and sparsely glan- dular-pubescent. Rosette leaves narrowly elliptic to elliptic or oblanceolate to narrowly obovate, short-acute, gradually narrowed to a short petiole or sessile, cuneate at the base, 6-18 cm long, 1.5-3.5 cm wide; cauline leaves oblong to elliptic or narrowly ovate to ovate, short-acute, truncate to subcordate at the base, sessile, 1.5-6 cm long, 1-3 cm wide; bracts oblong to broadly oblong or ovate, short-acute or subobtuse, truncate to subcordate at the base, sessile, those of the central and upper portions of the inflorescence much shorter than the capsule they subtend, 1-3 cm long, 1-3 cm wide; leaves mostly irregularly serrate, the teeth blunt or acute, plane or undulate along the margins; bracts usually red along the margins. Inflorescence branched. Floral tube (6.5-)8-10 cm long, often streaked and flecked with dark red. Buds narrowly oblong to lanceolate in outline, red at the junction of the sepals with the floral tube, 2-3.5 cm long, 5-11 mm thick. Sepals green to yellowish green, often streaked and flecked with red; apices of the sepals erect or divergent, 1-3 mm long. Petals very broadly obovate, yellow, often with a red spot at the base, 2-4 cm long. Anthers 7-13 mm long. Filaments 14-24 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, or the style long, the stigma held above the anthers at anthesis, 8-13 cm long. Stigma lobes 6-12 mm long. Ovary 1.7-2 cm long. Capsule mostly curved and with 4 clearly distinct cre- nate valves at the apex, 3-45 cm long, 3-4 mm thick. Seeds elliptic to broadly elliptic, brown, 1.5-2 mm long, 0.8-1.1 mm thick. Self-compatible; self-polli- nating and complex heterozygote. Gametic chromosome number, n — 7 (7 biva- lents, ring of 14 or ring of 12 and 1 bivalent at meiotic metaphase I). Flowering time: October-March. Type: Cultivated at Uppsala, Sweden, the sceds from the vicinity of Buenos Aires, Argentina, grown in 1752 or earlier, C. Linnaeus ( LINN, holotype, POM photograph). 510 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 FicunEs 154-155. Taxa of Oenothera sect. Oenothera subsect. Munzia.—154. O. grisea Chile, Valparaíso, Las Ventanas, Constance in 1965).—155. O. nocturna (Peru, Lima, San- tarius 2333). Distribution (Figs. 229-230): In Brazil only in Rio Grande do Sul; i Uruguay in the departments of Río Negro, Colonia, San José, and Montevideo; and in Argentina in the provinces of Corrientes, Entre Ríos, and Buenos Aires. — 1 Oenothera longiflora is unmistakable because of its dense long. villous pubes- cence and long floral tube. It is characteristic of sandy places along rivers and near the coast. This species is related to O. ravenii, as shown by similarities in habit, short bracts, reddish leaf margins, and the red basal spot on each petal. The seeds of O. longiflora are the largest in the series. Oenothera longiflora subsp. grandiflora is chromosomally homozygous, subsp. longiflora is chromosomally heterozygous. The latter has two complexes from subsp. grandiflora, but one of them appears to have been influenced by intro- gression from O. ravenii. Among the F, progeny derived from crossing the het- erozygote O. longiflora subsp. longiflora with O. ravenii, there are two classes of progeny, one intermediate and the other more similar to O. ravenii. The influence of the “pure” O. longiflora subsp. grandiflora genome upon the char- acteristics of O. longiflora subsp. longiflora is so strong, however, that it seems undesirable to recognize this particular complex heterozygote as a species sepa- rate from subsp. grandiflora. Oenothera longiflora subsp. grandiflora is evidently very rare and occurs 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 511 only at scattered localities. It may either have a relictual distribution, therefore, or be derived as a viable homozygous segregate from the chromosomally struc- turally heterozygous subsp. longiflora. KEY TO THE SUBSPECIES 1. Style long, the stigma held above the anthers at anthesis; buds 2.5- 3. E cm long; petals !!... ce SERRE $3948 a. subsp. grandiflora l'. Style short, the anthers prd a pollen directly € on the stigma at 4 buds em long; petals 2-3 cm long 17b. subsp. 8 17a. Oenothera longiflora subsp. grandiflora Dietrich, subsp. nov.—Fics. 50- 52, 128, 177, 213. Folia rosulae 2.5-3.5 cm lata; folia caulina 2-3 cm lata; bractea 1.5-2 cm lata, 1.5-2.5 cm longa. Gemmae 2.5-3.5 cm longae, 8-11 mm crassae; apices sepalorum divergentes, ca. 3 mm longi. Petala 3-4 cm longa, basi rubro-maculata. Nume ug gameticus chromosomaticus, n = 7; planta chromosomatice homozygotica. Rosette leaves 2.5-3.5 cm wide; cauline leaves 2-3 cm wide; bracts 1.5-2 cm wide, 1.5-2.5 cm long. Buds 2.5-3.5 cm long, 8-11 mm thick: apices of the sepals divergent, ca. 3 mm long. Petals 3-4 cm long, with a red spot at the base of each one. Anthers 10-13 mm long. Filaments 22-24 mm long. Stigma lobes 8-12 mm long. Style long, the stigma elevated above the anthers at anthesis. Self-compatible. Gametic chromosome number, n=7 (7 bivalents* at meiotic metaphase I). Type: Grown from seeds and cultivated in the Botanical Garden of Diissel- dorf, Germany, 19 Oct. 1972. Source: Argentina, Prov. Corrientes, Paso de los Libres, 27 Mar. 1964, A. Krapovickas & C. Cristóbal 11293 (MO-2155211, holo- type; CTES, DUSS, M, isotypes). Distribution (Fig. 229): Known only from the type locality and one locality in the province of Entre Ríos, Argentina. Specimen examined from cultivated plants ARGENTINA. CORRIENTES: Paso de los I dinde Krapovickas & Cristóbal 11293 (DUSS, M, Additional specimens examined: RGENTINA. ENTRE Rios: Dep Federación, Santa Ana at Río Uruguay, Burkart 29273, 26317 (SI). In 1962 and 1963 there appeared in Düsseldorf among the progeny of plants growing wild in the Botanical Garden of Buenos Aires (Gopel in 1961), one plant with 7 bivalents and another with a ring of 4 and 5 bivalents. Unfortu- nately, both of these strains were lost, since a test-progeny grown in 1973 yielded only plants with a ring of 12 and 1 bivalent which would therefore be referred to O. longiflora subsp. longiflora. l7b. Oenothera longiflora subsp. longiflora.—Fics. 55-57, 129, 214. Onagra 5 Moench, Meth. Pl. 1: 675. 1794. rype: The herbarium of Moench appar- ] longer exists. pues polymorpha H. Lév. race mn (L.) H. Lév., Mongr. Onoth. 364. 1909; Bull. cad. Int. Géogr. Bot. 19: 324. 1905 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ñ : b... * ^ E E * * a 2 a * * w Ficures 156-159. Taxa of Oenothera sect. Oenothera subsect. Munzia.—156. O. magel- lanica (Argentina, Mendoza, Santarius 1581).—157. O. villaricae (Chile, Cautin, Göpel in 1961).—158. O. hechtii (Argentina, Tucumán, Hecht 1964-81 ).—159. O. elongata (Bolivia, Tarija, Santarius 1956). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 513 O. polymorpha race longiflora var. sellowii Link & Otto ex H. Lév., Monogr. Onoth. 364. 1909; ull. Int. Géogr. Bot. 19: 324. 1909. LecrorypE: Cape Verde Islands, Brava, pide unc 900 m, 30 Mar. E ~ T. Lowe (BM; P, isolectotype). ox given in ng in Bolivia by H. eillé (anon moltsins L. var. aa (L. ) Hassler, Bull. Soc. Bot. Genëve, sér. 2, 5: 274. 1913. Sprague & Riley, Bull. Misc. Infor. 1921: 201. 1921. — Raimannia longiflora (L. Rosette leaves 1.5-2.5 cm wide; cauline leaves 1.5-2.5 cm wide; bracts 1-1.5 cm long, 1-1.5 em wide. Buds 2-2.5 cm long; apices of the sepals erect, 1-2 mm long. Petals 2-3 cm long, often with a red spot at the base of each one. Anthers 7-12 mm long. Filaments 14-22 mm long. Stigma lobes 6-8 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis. Gametic chromosome number, n = 7 (ring of 14* or ring of 12 and 1 bivalent** at mei- otic metaphase I). Distribution (Fig. 230): In Brazil only in Rio Grande do Sul; in Uruguay in the departments of Río Negro, Colonia, San José, and Montevideo; and in Argentina in the provinces of Entre Ríos and Buenos Aires. Not known from the province of Corrientes in Argentina, where O. longiflora subsp. grandiflora occurs. Specime ns examined from cultivated plants: JRUGUAY. COLONIA: Sandy places at the eos entrance of Juan L. Lacaze, Santarius 73*, UF TE, §1*, 82-84, 86,* 87, 88,* 89, 90 (DUSS; 81 also CTES; 73, 81 also M; 73, 81, 89 also MO). Dunes NW of fan L. on Santarius 109-115, 116* (DUSS; 116 sb. jn 109, Jh i MO). ARGENTINA, pe AiRES: Wild in the Botanical Garden of Buenos Aires, Göpel in 1961** (DUSS, M). La Plata, Hecht 1964-22* ( CTES, DUSS, M). Villa Ortuzor in Buenos Aires, Hecht 1964-123 (DUSS, N Additional specimens 3 RAZIL. RIO GRANDE DO SUL: oa Grande, Gaudichaud 1287 (P). URUGUAY. RÍO NEGRO: San Jav Puerto 8415 (MVFA); Herter 82853 (POM). coro- NIA: Colonia, Molfino 31-1650 (POM); N.N. 27 (BAA). Dunes near Riachuelo, Cabrera 3856, 3345 (LP). san JOSE: Arazati, Arrillaga 641 (MVFA). Playa Pascual, Arrillaga 741 (MV FA). MONTEVIDEO: Montevideo, Fruchard in 1874 (RSA); Barattini in 1939 N.N. (MPU); Lorentz 307 (G). Sta. Lucía, Fruchard in 1874 (US); Fruchard in 1875 Coast near Montevideo, Gibert 339 (K). Maluni, Rosa-Mato 1525 (LIL). Punta Gorda, Osten 4664 (G), 22709a (GH). Between La Colorada and Pajas Blancas, Lema 6458 (F, MVFA ). Miguelete, Berro 5340 Spee Cerro, Herter 162 (F, G, GH, HBG, LE, M, MO, POM, Z). Buceo, Felippone 2895, I (SI). Carrasco, Fruchard in 1875 (P Wo N 5784 (CORD, IS); Felippone 6097 (SI); pedis 343 (POM); Rosengurtt B418 (POM). Malvín, Felip- pone 3342 (SI). nocHa: Eastern coast, Tweedie 78 (BM ARGENTINA. BUENOS AIRES: Dock Sur, Hicken 784 (LIL, SI); Molfino 221 (POM). Pellegrini, Cabrera 6954 (LP, NY). Devoto, Hirschborn 40 (POM). Maciel, Molfino 25842 5 La Plata, Hirschhorn 676 (POM). Paternal, Parodi 8993 (GH, K, POM). Caron, Franqueville (P). ENTRE RIOS: w- del Uruguay, Lorentz 1230 ( CORD, G, SI). Con- cordia, Spegazzini S8 (BAB). Santa Ana, Gamerro 1310 (LP). Berduc, Crovetto d» Piccinini 4654 (BAB). El Palmar, Pedelaborde in ” 1940 (SI). Between Gómez and Emb. Ferrari near Concordia, Meyer 10974 Spec cimens from outside of Séuth America (naturalized ): Near Bayonne, Lange in 1851 (P); Bordére e: 1877 (G); Blanchet in 1880 (P). Between Bayonne and Boucau, 1881, Blanchet 70 (LISU, LY, MPU, P); Neyraut in 1889 (COI, MPU); 1933, Hibon 1461-2 (P). Boucau, Deflers in 1879 (MPU); Neyraut in 1902 (LY); Willkomm (COD. Dep. Landes, Riviére, ED gud in 1881, in 1882 (LY). Biar- ritz, Henry in 1903 (B). Anglet, Neyraut in 1905 (MPU ). PORTUGAL. Acóres, Fayal, Donat in 1868 (G, P). MADEIRA: Serro de S. Roque, 1865- ~ - — "am! — — "d w 514 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 1866 SPAIN. CANARY ISLANDs: Puerto de La Orotava, N.N. 2352 (K). SOUTH AFRICA. CAPE: Camps Bay, BS. in 1846 (K); Prior in 1846 (PRE). Wellington, Thomson in 1881 (PRE). Cape t Good Hope, Pappe (K). USTRALIA. NEW SOUTH WALES: Raymond Terrace, Coans in 1961 (NSW-65372). Prin- cess Highway at crossing of Minimurra River, 1 mi S of Albion Park, Raven et al. 25895 in 1970 (MO, NSW); Briggs 3950 in 1970 (NSW). Hybrids with O. stricta subsp. stricta have been collected in France, where both species grow together near Biarritz: Biarritz, Connan in 1907 (MPU). Anglet, 1947, Jallu 5257 (LD), 1964, 7832 (LISE). Early specimens from plants cultivated in Botanical Garden Erlangen, Germany, Herb. Schreber in 1779 (M). Paris, in 1781 (MPU). Erlangen, Herb. Schreber à in 1800 (M). Halle, Germany, Fischer in 1801 (L). Montpellier, France, in 1808 (MPU). Hamburg, Germany, Hun seeds of the Botanical Garden Kiel, in 1815 (HBG). Munich, iu los Herb. Zuccarini in 1835 (M; as O. nervosa). Paris, Abbé Pouret in 1847 (P). Hort. Cantab., seeds from the Canary Islands, Herb. Gray in 1867 (GH; as O. canariensis). Oenothera longiflora 5 in literature from outside of South America: France: Gandoger (1886: 49); Rouy & Camus (1901: 201); Coste (1903: 81); Thel- lung (1912); Bonnier (1921: 29); Fournier (1937: 598); Issler, et al. (1965: 357); Raven (1968). BELcivuM: Jean (1975). PorrucaL: Gandoger (1886: 49); Coutinho (1913: 427; 1939: 508 AzonEs: Trelease (1897: 114). MADEIRA: Lowe (1868: 275); Menezes (1914: 71). SpAIN: Raven (1968). Sourn Arnica: Phillips (1917: 99; as O. villosa); Adamson & Salter (1950: 606). Jamaica: Grisebach (1860: 273). 1866, Mandon 441 (G, P). Mountains above the Povoação, N.N. 766, 768 (K); Lowe in (BM). — ; Raven (1968). 18. Oenothera catharinensis Cambess., in St.-Hilaire, Fl. Bras. Mer. 2: 270. 1829.—Fics. 3, 53-54, 130. O. mollissima sensu Munz, Amer. J. Bot. 22: 659. 1935, pro parte. Erect annual herb, not forming a rosette, bushily branched near the base, 1-5 dm tall. Plants densely to sparsely strigillose and villous in the lower por- tions, densely to sparsely long-villous and thickly to sparsely glandular-pubes- cent elsewhere. Cauline leaves cultrate to narrowly lanceolate, acute, acute to rounded at the base, sessile, 3-5 cm long, 0.7-1 cm wide; bracts narrowly oblong to lanceolate, acute to rounded at the base, sessile, 2-4 cm long, 0.5-0.8 cm wide, mostly shorter than the capsules they subtend; leaves plane and irreg- ularly serrate with blunt teeth. Inflorescence branched. Floral tube 3-4 cm long. Buds lanceolate in outline, 1.8-2.2 cm long, 5-6 mm thick, green or yel- lowish green; apices of the sepals erect, ca. 1.5 mm long. Petals very broadly obovate, retuse, 3-3.5 cm long. Anthers 7-10 mm long. Filaments 15-18 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 4—5.5 cm long. Stigma lobes 5-8 mm long. Ovary 1-1.5 cm long. Capsule 3-4 cm long, 3-4 mm thick. Seeds elliptic in outline, 1.5-1.8 mm long, 0.6-0.8 mm wide, brown. Gametic chromosome number, n = 7 (7 bivalents*, ring of 14** or ring of 4 and 5 bivalents*** at meiotic metaphase I). Flowering time: October-March. Type: Brazil, Santa Catarina, Îlha Santa Catarina, Apr. (1816-1821), Auguste de St.-Hilaire 1721 (P, holotype, F and GH photographs; MPU, P, isotypes). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 515 Distribution (Fig. 229): Apparently only along the coast of the state of Santa Catarina, Brazil. Specimens examined from cultivated plants: BRAZIL. SANTA CATARINA: Coast near Itapema, Hatschbach in 1971** (DUSS, M, MO). Isle of Santa Catarina, Praias Inglese, Conrad 5 Dietrich 8***, 9, 10, 13* (DUSS). bon nal specimens examine L. SANTA CATARINA: Sania Catarina, N.N. (GOET). Isle of Santa Catarina, Gaudi- chaud 259 (P). Isle of Santa Catarina, Pantano do Sul, Klein et al. 5824 (HBR, RSA). Rio Vermelho, Klein et al. 5801 (HBR, RSA); Perdonnet 233 (G). Laguna, Reitz & Klein 185 (RSA), 137 (HBR, RSA Oenothera catharinensis has generally been regarded as a synonym of the complex structural heterozygote O. mollissima (Munz, 1935), but it can be dis- tinguished from that species by its strigillose pubescence, which is never present in O. mollissima, and its larger flowers. The pubescence, flower size, and short bracts of O. catharinensis indicate that it is a relative of O. ravenii rather than of O. mollissima. I have been unable to discover any evidence that O. catharinensis has par- ticipated in the origin of any complex heterozygote. From this, I conclude that it may be a species of relatively recent origin derived from an ancestor similar to O. ravenii as an obligate annual strain with altered characteristics (Fig. 8). The occurrence of complex heterozygote strains with a ring of 14 chromosomes may reflect a situation similar to that in O. affinis and O. odorata. 19. Oenothera indecora Cambess. in St.-Hilaire, Fl. Bras. Mer. 2: 268. 1729.— Fics. 2, 44-46, 131-132, 178-179, 215-217. Erect annual, forming a rosette, unbranched or with a moderately or strongly branched main stem and arcuate to obliquely ascending side branches arising from the rosette, 2-6 dm tall. Entire plant sparsely long-villous with erect hairs, densely to sparsely short-villous, and densely glandular-pubescent; or only densely short-villous and densely glandular-pubescent. Rosette leaves narrowly oblanceolate, acute, gradually or + abruptly narrowed to the petiole, 4-8 cm long, 0.2-1.3 cm wide; cauline leaves very narrowly elliptic to lanceolate, acute, acute at the base, sessile, 2.5-7 cm long, 0.2-1.2 cm wide; bracts very nar- rowly elliptic to elliptic, acute, acute at the base, sessile, longer than or about the same length as the capsules they subtend, 1.5-5 cm long, 0.2-1 cm wide; leaves plane or undulate at the margins, irregularly serrate with blunt teeth. Inflorescence branched. Floral tube 0.5-1.5 cm long. Buds oblong to broadly elliptic in outline, 0.2-0.8 cm long, 1.5-4 mm thick. Sepals green or yellowish green, often = densely flecked with reddish brown; apices of the sepals 0.5-1 mm long, erect. Petals broadly elliptic to very broadly obovate, yellow or bright yellow, 4-10 mm long. Anthers 1.5-4 mm long. Filaments 1.5- 7 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 1-2 cm long. Stigma lobes 1-2 mm long. Ovary 1-1.5 cm long. Cap- sule 2-3 cm long, 1.5-2 mm thick. Seeds broadly elliptic to rotund in outline, 0.7-1.3 mm long, 0.3-0.5 mm thick. Cleistogamous, almost always fertilized in bud. Gametic chromosome number, n = 7 (7 bivalents or ring of 14 at meiotic metaphase I). ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficungs 160-163. Taxa of Oenothera sect. Oenothera subsect. Munzia.—160. O. pseu- doelongata (Bolivia, Cochabamba, S ius 1988).—161. O. cordobensis (Argentina, Cór- doba, Gópel in 1961).—162. O. siambonensis (Argentina, Tucumán, Göpel in 1961 ).—163. O. brevipetala (Bolivia, Cochabamba, Santarius 1972). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 517 Type: Brazil, Rio Grande do Sul, sandy places near the farm Manguiera, not far from the city of Rio Grande, Aug. (1816-1821), Auguste de St. Hilaire 1872 bis (P, holotype, F and GH photographs; MPU, P, isotypes). Distribution (Fig. 231, 233): In Brazil from the states of São Paulo to Rio Grande do Sul; in Uruguay in the departments of Cerro Largo, Treinta y Tres, Rocha, Maldonado, Montevideo, San José, Artigas, Riviera, Salto, and Rio Negro; in Argentina throughout the northern and eastern portions of the repub- lic, in the provinces of Jujuy, Salta, Tucumán, Catamarca, La Rioja, Formosa, Chaco, Santiago del Estero, Santa Fé, Córdoba, San Luis, Misiones, Corrientes, Entre Rios, and Buenos Aires; in central Paraguay; and in Bolivia in the vicinity of La Paz (subsp. boliviensis). Together with O. mendocinensis and O. verrucosa, O. indecora contradicts the rule that chromosomally homozygous entities in the genus are usually large- flowered. In comparison with O. mendocinensis, O. indecora has wider leaves, shorter capsules, smaller seeds, and never any strigillose pubescence. KEY TO THE SUBSPECIES 1. Plants soir: short villous and glandular-pubescent, appearing glabrous to the naked e 2. Bracts 0.5- 1.3 cm wide; buds 4-8 mm long; petals 0.5-1 cm E — . 19b. Qd bonariensis 2’. Bracts 1.5-2 mm wide; buds 2-3 mm long; petals 0.2-0.3 cm E ° CTC e a IR RE VNB subs sp. _boliviensis ]”. Plants ‘sparsely long villous, densely short “gildas, and densely be "a apay. pubescent to the naked eye; bracts 0.5-1 cm wide; buds 3-8 mm m s 0.4-1 cm long — 6 : — 19a. subsp. ri ER 19a. Oenothera indecora subsp. indecora.—Fics. 131, 178, 215-216. Onothera polymorpha H. Lév. race mollissima (L.) H. Lév. var. "d ( Cambess.) H. Lév., Monogr. Onoth. 365. 1909; Bull. Acad. Int. Géogr. Bot. 19: Raimannia indecora (Cambes s) Sprague & Riley, Bull. Misc. Infor. T5 201 1921. Oenothera argentinae H. Lév. & Thell. var. “aqp Kloos & Thell., Ned. Kruidk. A rch. 1921: 0. LECTOTYPE: ub N t, oatfield near he Karelke meal p aien 13 Aug. 1920, A. W. Kloos ( L-95264486; bye isolectotype Plants 2-4 dm tall. Entire plant sparsely long-villous, densely short-villous, and densely glandular-pubescent. Rosette leaves and cauline leaves 5-7 cm long, 0.5-1 cm wide; bracts 3-5 cm long, 0.5-1 cm wide. Buds 0.3-0.8 cm long, 2.5-4 mm thick. Sepals = densely flecked with reddish brown, seldom immaculate. Petals 0.4-1 cm long. Anthers 2-3 mm long. Filaments 3-6 mm long. Seeds elliptic in outline, 1-1.3 mm long, 0.4-0.5 mm thick. Cleistogamous. Gametic chromosome number, n — 7 (7 bivalents* at meiotic metaphase I). Flowering time: Brazil, September-July; Uruguay, Paraguay and Argentina, September- Distribution (Fig. 231): In Brazil from the states of São Paulo to Rio Grande do Sul; prevalent in the coastal departments of Cerro Largo, Treinta y Tres, Rocha, Maldonado, Montevideo, Colonia, Canelones and San José in Uru- guay, and also in the departments of Ártigas, Salto, Lavalleja and Rivera; in Argentina known from the provinces of Misiones, Corrientes, Entre Ríos, For- 518 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 mosa, Chaco and Córdoba; in Paraguay only from Encarnación in the southeast of the country. Specimens examined from cultivated plants: RUGUAY. MALDONADO: San laces in tha southern part of Piriapolis, Santarius 147, 150*, 151, 152, 153*, 155, 156 (DUSS; 150 also CTES; 150, 152 also M, MO). FLORIDA: At the railroad 1-3 km NE and SW of Mansavillagra, Santarius 205*, 20 5655 208* 209“, 212“, 213* (DUSS; 205, 208, 209 au CTES, M; 205, 206, 208, 209 also MO). Lavadero San Pedro, Hecht 1964- 63* (DUSS, M, MO). itional specimens examine BRAZIL. SAO PAULO: Pinheiros, Gehrt 34390 (POM, SP). PARANA: Curitiba, Hoehne 23080 (POM, SP). Porto Vitória near Curitiba, „ = 1969 (UC). SANTA CATARINA: Itajai, Klein 2567, 2686 (HBR, RSA); Mueller s.n. (R), 397 (K). Near Palmitos, 400 m, Smith & Reitz 12584 (HBR, US). Rio Irani 12 km S of PUN des Guedes, 500-600 m, Smith & Klein 12903 (HBR, RSA). Rio Uruguai near Itapiranga, 200 m, Smith & Klein 13184 (HBR, RSA). Ilha de Santa Catarina, Klein & Bresolin 8338 (HBR). Araranguá near Sombrio, Reitz C748 (BR, HBR). Pirão Frio near Sombrio, 10 m, Reitz & Klein 9072 HE Est. Exp. de Videira, Santos 2894 (R). Tubarão near Laguna, Ule 1256 (HBG, P). ANDE DO SUL: Near Trés de Maio, Santos 2763 (R). Serro de Pustral near 0 Maria. "Vidal 01199 (R). Palmeira near Xingu, 650 m, Bornmiiller 745 (GH). Sao Sep r Cacapava, 5 8734 (MVFA). City of Rio Grande, Malme 270 (S). Viera, eee 36795 (SP), 4301 (BR, GH, NY, POM, US). Between Pelotas and Rio Grande, Krapov vickas d» Cristóbal 22901 (MO). Pelotas, da Costa Sacco 1116 (F, HBR); Casagrand et al. 13 (CTES). Facenda Saledade near Rio Pardo, Jürgens 25, 97 (B); Vidal 01526, 01533 (R). Montenegro near L. S. Pedro, Seh- nem 3833 (B, SI). San 5 Reuter in 1953 (R); Eugenio 201, 206 (NY); Rambo 293 LIL). Uruguayana, Palacios & Cuezzo 202, 227 (LIL); Vidal 01315, 01318 (R). Passo de Ricarde at Rio Piratini, Poraba 6773 ( RB). Toca do Tigre near Itapoan, Rambo 48931 (LIL, MO), 48934 (HBR, RSA). Porto Alegre, Rambo 293 (SP). Gloria near Porto Alegre, Rambo 978 (SP), 27196 (LIL, S), 28995 (RSA, LIL). Villa Manresa near Porto Alegre, Rar nbo 37466 (F, HBR, LIL, RSA), 57073 (HBR), 55570 (B, HBR). Balneario Ipanema near Porto Alegre, Emrich 1126 (LIL). São Joño near Pôrto Alegre, Reineck & Czermak 30 (HBG). Praia Tramandahy near Porto Alegre, Vidal in 1913 (R). Montenegro, Rambo 29702 (LIL). Pareci near Montenegro, Rambo 29702, 42445 (LIL). Amaral Ribeiro near Taquara, Rambo 42391 (F, LIL). Travessáo near Hamburgo, Rambo 42181 (LIL). Osorio, Rambo 48887 (LIL). Morro Sapucaia near Pérto Alegre, Rambo 37381 (SI). Est. Azevedo, Rambo 43276 (LIL). URUGUAY. ÁRTIGAs: Cuareim, Berro 2319 (MVFA). Santa Amaro, Jürgens 168 (B). SALTO: Dayman, Osten 5258 (SI, US). RIVERA: pecie 200-230 m, Wright in 1928 (BM). FLORIDA: Mansavillagra, Rosengurtt B790 (POM). san josé: Sierra Mahoma, Izaguirre 2591 (MVFA). Rincón del Pino, Izaguirre 9543 (MVFA). COLONIA: Between E and Monte- video, Burkart 29012 (SI). 5 Carrasco, Rosengurtt 861 (POM); Berro 7550 (MVFA). Sta. Lucía, Fruchard 788 (P). cerro Largo: Est. Perdomo near Ruta 8 and 5 5 Arrillaga 2358 (F). pes een Río Negro and Acegua, Rosengurtt 85la (POM). REIN s: Km 323 at Ruta 8, Marchesi 2268 (MVFA). LAvALLEJA: La Lorenzita, 5 9132 (MVFA). Abra de Perdano, Olano 8838 (MVFA). Cerro Arequita near inas, Krapovickas & Cristóbal 16149 (CTES). CANELONES: Sta. Lucia, Gibert 988 (K); Felippone 5574. 5589 (SI). nocHA: La Coronilla, Brescia 3984 (MVFA). Parque Sta. Tereza, Rodriguez 8146 (MVFA). MALDONADO: Sierra de las Animas, Izaguirre 10751 (MVFA ). H S H, UC, PARAGUAY. Encarnación, Hassler 1476 (SI); Bertoni 4532 (LIL). ARGENTINA. MISIONES: Bonpland, Van de Venne 275 (POM). p Schinini igpay (CTES). connENTES: Dep. Tah at Río Miriñay near Ruta 23, Schinini et al. (CTES, MO). ENTRE Rios: Concepción del Uruguay, Lorentz 424 | pro parte (GOET); 5 art 28762 (MO). Between Colón and Concepción, Burkart 28991 (MO). San Salvado, Baez 52972 (BAB). Arroyo Maimol near Colón, Nicora 3238 (LIL). Km 249 at Ruta 14 near Con- crodia, Burkart 22664 (MO). Arroyo Isletas, Burkart 27034 (MO). a ao 23759 (MO); Anetto in 1891-92 (CORD). Gualeguaychi, Meyer 10331 (LIL). danos, Burkart 3470 (POM). Rio Tale, Schulz 284 (LIL). Paranacito, Ragonese 6 ere FORMOSA: N.N. in 1904 (SI-4860). Reventón near Pilcomayo, Morel 1913 (LIL). caco: Dep. Independen- cia, Campo Largo 8 km from J. Mármol, Bacigalupo et al. 9601 (BAA, MO, SI), 9604 (BAA). Puerto El Colorado, Rojas 12113 (LIL). Dep. Napalpi, C. del Bermejo, Burotorich 535 (LIL). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 519 CÓRDOBA: Vicinity of Alta Gracia, Hunziker 7710 (CORD). Sierra Ochoa near Sta. María, Stuckert 13562 (CORD). Between Est. Burmeister bn Onagoity near Gral. S. Martín, Di Fulvio 21411 (CORD). At km 571 of Ruta 35 between Huinca Renancó and Río Quinto, Hunziker & Di Fulvio 21430 (CORD). Near reece " Bundle 6951 (CORD). Specimens from outside of South America: NETHERLANDS. Wormerveer, meal factory, Jansen & Kloos in 1913 (BAS). France. Dép. Nord, Dunkerque, Bouly de Lesdain in 1924 (BAS). HUNGARY. Györ at river Danube, 1916, Polgar 2468 (BAS). Plants intermediate between O. indecora subsp. indecora and O. indecora subsp. bonariensis are occasional in the zone of overlap between these two enti- ties and may be recognized by the fact that they are only sparsely long villous. All plants of this species are in effect cleistogamous, fertilization taking place in bud well before the unfolding of the flowers; and this factor alone must drasti- cally limit the possibility of hybridization and thus help to maintain the integ- rity of the two subspecies where they come together. One of the genomes of the complex heterozygote O. montevidensis is derived from O. indecora subsp. indecora. 19b. Oenothera indecora subsp. bonariensis Dietrich, subsp. nov.—Fics. 2, 44-46, 132, 179, 217 Oenothera S Mr ia Torr. & A. Gray f. erecta Thell. & Zimmerm., Repert. Spec. Nov. Regni 'g . 1916. TYPE: Germany, port of Ludwigshafen, Sep. 1909, F. Zimmermann — argentinae H. Lév. & Thell. ex H. Lév., Monde Pl., sér. 2, 18 (108): 52. 1917; H. Thell., Repert. Spec. Nov. Regni Veg. 15: 133. 1918. LECTOTYPE: Germany, riu of Here en at the Rhine near an oil refinery, 26 Sep. 1915, Bonte (Z; BA, POM, isolectotypes). Léveillés mention of Onothera argentinae in Monde Pl., sér. 2, 18 (108): 52. 1917, is not a nomen nudum, as stated by Lauener (1972: 424). Ned. Kruidk. Arch. 1921: 100. 1921. LECTOTYPE: Netherlands, Rotterdam, Mao haven pese factory, 10 Sep. 1920, A. W. Kloos (L- 954264523; BAS, isolectotype . argentinae var. brevipila Kloos & Thell., Ned. Kruidk. Arch. 1921: 100. 1921. LECTOTYPE: etherland, Weert, oatfield near the Karelke meal factory, 13 Aug. 1920, A. W. Kloos (L- 954264489; wel Eei iar ). . indecora sensu Mu hysis 11: 281. 1933, pro parte; Amer. J. Bot. 22: 658. 1935, pro arte; sensu Munz, "i Brasílica 9(41): 53. 1947, pro parte. . argentinae Erlangen? Haustein, Z. Indukt. Abstammungs- Vererbungsl. 84: 418. 1952. . indecora "Argentinae," "Buenos Aires" and "Reconquista" Cleland, Jap. J. Genet. 43: 332. 1968. o ° a = ya m = ~ š a eS < £e — + < — a. ° a 2 ° ° ua D - 2 e RF o coo Plantae 2-6 dm altae, ubique dense breviter ber denseque glanduloso-pubescentes, sine lente ut videtur glabrae. Folia rosulae 5-7 cm longa, 0.5-1.3 cm lata; bractea 3-5 cm longa, 0.5-1.3 cm lata. Gemmae 0.4-0.8 cm longae, 2.5—4 mm crassae. Sepala viridia vel flavo- virentia, raro rubro. Semina ambito late elliptica, 0.7— E mm longa, 0.4—0.5 mm crassa. Nu- merus gameticus chromosomaticus, n — 7; planta chromosomatice homozygotica vel heterozy- gotica complexa. Plants 2-6 dm tall. Entire plant densely short-villous and densely glandular- pubescent, apparently glabrous when viewed without a lens. Rosette leaves 5-7 cm long, 0.5-1.3 cm wide; bracts 3-5 cm long, 0.5-1.3 cm wide. Buds 0.4-0.8 cm long, 2.5-4 mm thick. Sepals green or yellowish green, rarely flushed with red. Petals 0.5-1 cm long. Anthers 1.5-4 mm long. Filaments 4-7 mm long. Seeds broadly elliptic in outline, 0.7-1 mm long, 0.4-0.5 mm thick. Self-polli- 520 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ficures 164-165. Taxa of Oenothera sect. Oenothera subsect. Munzia.—164. O. acuti- arpa (Argentina, Tucumán, Gópel in 1961).—165. O. tucumanensis ( Argentina, Tucumán, Santarius 1657) nating and often cleistogamous. Gametic chromosome number, n = 7 (7 biva- lents* or ring of 14** at meiotic metaphase I). Flowering time: September-July. Type: Argentina, Prov. Buenos Aires, Isla Santiago near La Plata, 24 Nov. 1935, A. L. Cabrera 3406 (NY, holotype; F, G, LIL, LP, POM, SI, UC, isotypes). Distribution (Fig. 233): In Brazil from the state of Sào Paulo to Rio Grande do Sul; in Uruguay the stations are all in the western and southwestern depart- ments of Río Negro, San José, and Florida; in central Paraguay and throughout the range of the species in Argentina. Specimens examined from cultivated plants: ARGENTINA. BUENCS AIRES: On sandy soil 2 Isla Santiago near La Pl: ata, Santarius 275*, 276* 277*, 278*, 281*, 284*, 285, 287, 291*, 295, 297, 299 (DUSS; 275 also CTES, M, SP; 275, 276, 277, 278, 284 also MO). Buenos yis Cle land 1967-508* 8925 DUSS, MO). TUCUMÁN: Rio Sali near Tucumán, “El Cadillal," Göpel in 1961* (DUSS, MO). CORRIENTES: Santo Nani Krapovickas 16404 ( DUSS, MO UAY. COLONIA: Sandy = at Toads in western part of Juan L. Lacaze, Santarius 68**, 72 (DUSS; 68 also M, MC ARANÁ: Corredeira Paulista at Rio Jaguariaiva near São José da Ba Paulista, Hatschbach 25564 (DUSS). ATED: O. argentinae from Erlangen, Germany, received 1960* (CTES, DUSS, M, MO). Representative specimens examined: BRAZIL: SAO PAULO: Alto da Serra, Gehrt 39948 (POM, SP). paraná: Curitiba, Dusén 13292 (F, MO, S), 2219 (R), 15827 (GH, LE, NY, S). Jaguariaiva Dusén 13189 (S, SI). SANTA CATARINA: Itajai, Klein 6859 (HBR, RSA). Nova Teutonia, Ploumann 583 (RB). nio ~ 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 521 RANDE DO SUL: RE Soni Vidal 01328 (R). Rio Grande, Malme 110 (S). Rio Pardo, * in 1923 CES Pórto Alegre, Lindmann 363 (S). GUAY. RÍO NEGRO: 98 Mercedes, isles of Rio Negro, Rosengurtt 855 (POM). sAN JOSE: Rio Santa bu Rosengurtt 859 (POM). FLORIDA: La Palma, Herter 2105 (F, MO, GUAY. 1 „ 4117 (C, F, MO, NY, S, SI, US). Villa Elisa near Asun- ción, 5 5118 (C tanical m of Asunción, Rojas 3174 (POM). Asunción, Ba- lansa 2219 (K, P EN yi de Maracay near river Ca apibary, Hassler 4409 ( BM, K, NY, P, UC, W). Sma, Trinidad, aap i in 1913 (Z). Without exact locality, Bonpland in 1833 (P). ARGENTINA. BUENOS AIRES igre near Garin, Lanfranchi 485 (LP). El Socorro near Pergamino, Parodi 7401 (CH). Campana, Krapovickas 2593 (BAB, d Hunziker 1044 (CORD, LIL, RSA). Juinez, Molfino 146 (POM). Est. Las Palmas near Port. Zárate, Boelcke 13152 (BAA). Villa Ortuzor, Parodi 9900 (K, POM, US). Isla Paulino, Cabrera 7385 (F, CH). Ramalla at Río Paraná, Burkart 12767 (SI). Pavón at Rio Paraná, Pennington 171 (CORD). Isla Maciel, Krapovickas 222 (LIL). Federación, Birabén 5127 (LP). MISIONES: Villa Venecia near Lesudro N. Alem, Krapovickas & Cristóbal 15970 (BAB, CTES, MO) Deseado near Frontera, Pierotti 5248 (LIL). 20 km S of Bernardo de Irigoyen, 150 m, Kra- povickas & Cristóbal 13714 (BAB, C, CTES, UC). Puerto Polana near Cainguas, Schwarz 7920 (LIL). Corpus, Scala 257 (LIL). Santa Ana, Rodriguez 647 (GH, LIL, POM, SI, UC). Orera near Candelaria, Pierotti 5265 (LIL). Loreto near Candelaria, Montes 14665 (NY). L, S). Puerto Cazador near San Ignacio, Posadas, Grondona & Spegazaini 1111 (BAB). Puerto istuele near Iguazu, Montes 9291 (GH, LIL). Eldorado near Iguazú, Schmidt 2097 (LIL). Apóstoles, Krapovickas et al. 15485 (CTES). corrmnres: Paso de la Patria near San Cosmé, Tressens et al. 119 (BAA, CTES, F, LP, by 25 Ibarrola 775 (LIL, NY). San Luis de Palmar, Ibarrola 3260 (LIL). Laguna Rincón near Gral. Paz, Schwarz 8554 (LIL). Loma Alta near Ituzaingó, Pierotti 600 ALILI. Between Río Empedrado and Ruta 12, Quarín e Schinini 1297 (CTES, Nn ^ = € a a N A e ° S r^ — E^ Pedersen 117 (P, S, US). Santa x Parodi 12061 (BAA). Dep. Capital, between Arroyo Riachuelo and Ruta 12, Krapovickas 2 Cristóbal 13559 (CTES, MO). Tabay near Concepción, Krapovickas & Cristóbal 11709 (CTES, K, US). Loreto near San Miguel, Mroginski 42 (CTES). Laguna Mansa near Paso de los Libres, Schinini 7603 (CTES, MO). ENTRE Rios: Paraná, Boelcke 1268 (BAA). Isla del Pillo near Victoria, Burkart 8735 (F, POM). Colón, Burkart 24874 (MO). Ibicuycito, Burkart 27309 (MO). San Carlos near Concordia, Crovetto & Grondona 4139 (BAB). Delta del Paraná, Arroyo Martinez, Boelcke 901 (BAA). Federación, Meyer 11131 (LIL). Formosa: Colonia d e N.N. 98 (BAB). caco: Margarita Belén, Aguilar 1078 (LIL); 77 7 (CORD). Las Palmas, Jörgensen 2489 pro parte (POM). Fontana, Meyer 455 (LIL, POM, SI). Vis-a-vis of Corrientes, Alboff in 1890 (NY). Colonia Benitez, Schulz 502 r. 10075 (BAB). SANTA FÉ: Mocovt, Venturi 94 ( BAB, CORD, LIL, P, POM, SI). E “suq Nu 3389 (LIL, RSA). Esperanza near Las Colonias, Huidobro 3263 (LIL, RSA). C a Mascias near 0 aray, Pueyo 105 (CTES). Rosario, Burk- art 8772 25 SI). Laguna ates pie 807 (L i 1 I San Justo, Erbaggi & Loin 966 (SI). i Cristóbal, Balegno 593 (LIL). Recon- quista, Parodi 11129 (POM), 11146 (BAA, POM); Burkart 5795 (F, SI). SANTIAGO DEL Es- TERO: Colonia Jaimez near Robles, Luna 1329 (LIL); Ruiz in 1948 (RSA). Zurena near Robles, d 441 (LP). La Isleta between Ceres and Colonia near Rivadavia, Hunziker eda. (CORD). Selva near Rivadavia, Balegno 462 (LIL). Parque Aguirre in Santiago del o, Arganaraz 440 (LIL). El Puestilo near Guasayán, Cuezzo 2405 (LIL). La Banda, O"Donell 4248 (LIL). corposa: Hieronymus in 1878 (BR, FR, K, LE, P). San Vicente, Kurtz 576, 577 (CORD). Río Primero, Kurtz 2625 (CORD). Capilla del Monte, Hossens 364 (CORD). Emp. Tanti near Punilla, De la Sota 3075 (LIL). Río Segundo, Sublis 1080 (CORD). Ascasubi near Tercero Arriba, Krapovickas 6391 (BAB, CORD). Cárcano near San Martín, 5 7403 (LIL). Casa Bamba near Colón, Stuckert 23732 (CORD). Near Villa Do , Hunziker 13204 (RSA). Alta Gracia, Rande 6761 (CORD, LIL). Est. La Redu- ción in che 8 Sierra Chica, Pastore 348 (SI). san Luis: Burkart 12086 (SI). juyuy: Agua Negra near Ledesma, Fabris et al. 3044 (LP). Villa Achaval, a 20059 (LP). SALTA: Río San Francisco near Orán, Rodríguez 1155 (GH, LIL, POM, SI). Puerta Verde near Santa 2° sección, N.N. 603 (LIL-216719). Vallecito near Metán, Panes 384 (LIL). TUCUMÁN: Cerro San Javier, Boelcke 2911 (BAA, MO). Rio Sali, Venturi 898 (BAA, GH, LIL, POM, SI, US). Chanar Pogo near Leales, Venturi 491 (LIL, LP, MO, POM, SI). Agua Dulce near Ne — 5 ° 2 (de Qu D m E E. 2 O~ 2 d — ° 8 > D p = = © — O 599 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 64 Leales, Monetti 1163 (GH, LIL). Río Colorado near Famaillá, Carenzo 3425, 2429 (LIL). Yerba Buena, Meyer in 1949 (LIL). El Morado PES Burroyacü, Peirano 10085 (LIL). cATA- MARCA: Andalgala, Jórgensen 1054 pro parte (LIL). Choya near Andalgala, Castillon 14417 ( , LIL). San Antonio, Castillon 1283 (LIL). Río del Valle near Piedra blanca, Lillo 9085 (LIL). LA RIOJA: Quebrada de Soria, 8 km SE of d 5 16638 (CORD). No- nogasta, Castellanos 27-2011 (POM). F. s saw: oe 6 (LIL). Dique de Alta, Hun- ziker " Fabris 14441 (CORD). san juan: Valle Fertil, Hunziker 16698 ( CORD). Specimens from outside of South America viles ize Germany. Humboldt mill near Tegel, Berlin, Schulz in 1896 (B). Port of Ludwigshafen, Zimmermann in 1907 (L). Port of Neuss, 5 in 1926 (BAS). Finkenwerder near Ham- burg, Christiansen in 1926 (BAS). Emmerich, 1930, Kern & Reichgelt 12180 (L). NETHERLANDS. Wormerveer, 1913, Kloos 57 (L), in 1916 (L), in 1923 (L). Rotterdam, 1904, m d Wachter 13287 (L), in 1920 (L); Kloos in 1920 (BAS, L). IC . Nord, Dunkerque, 1926, Bouly de Lesdain 585 (BAS). 5 TUGAL. Barreiro near Estremadura, 1958, Rainha 3703 (LISE). Moita, 1961, Rainha 4844 (LISE). RuovesiA. Distr. Selukwe, Ferny Creek, ca. 1,190 m, p Wild 4981 (K, MO, SRGH); 1966, 5 1498 (SRGH). Distr. Gwelo, 1,280 m, 1965, Biegel 475 (SRGH). ANA, Distr. an Eastern, Gaberone, Content Farm, 1972, Kelaole A 78 (SRGH). So OUTH UCA. TRANSVAAL: Near Pretoria, 1930, Moss 18261 (BM). Distr. Letaba, 1958, Scheepers 270 (BM, K. MO, PRE, SRGH). Duiwelskloof, 945 m, 1958, 5 270 (K). Distr. Wolmaranstad, in 1963 (PRE- 29488). Pta. Harde near Eloffsda l, 1963, Hanekom 1663 (K, SRGH). NATAL: Distr. Ifafa River, ca. 610 m, 1948, Gerstner 6932 (PRE). Distr. Port Shepstone, St. Michaels-on-Sea, 905 Strey 7094 (K). Distr. Umzinto, Shelley beach, 1967, Strey 7284 (K). ORANJE FREE STATE: Bloemfontein, 1956, Gemmell 6737 (K). Distr. Swinburne, Rensburgshop, 1961, Jac 15 91 (PRE). Distr. Kimberley, ca. 1,220 m, 1961, Leistner 2931 (K, PRE). Distr. Frankfort, Strydow (K, PRE). rLEsoruo: Maseru, 1969, Wil- liamson 22 (K, SRGH). cape: Hout Bay near Rondebosch, 1932, Adamson 2198 (BM). Cape 5 1933, Salter 4001 (K). Kuils River, near N 1953, Parker 4849 (K). Stellen- sch, 1952, Parker 4839 (K); 1964, Taylor 5643 OUTH WEST a. Windhoek mountains, 9 near Finkenstein, 2,000 m, 1965, Seydel 4223 (MO AN Da e Stableford in 1953-54 (K). AUSTRALIA. NEW SOUTH WALES: Birdwood near Yarras, Noonen in 1961 ( NSW-135161 ). “Carp View" near Gilgandra, Holawich in 1965 (NSW-135156). La Peraouse, Coveny in 1965 (NSW), in 1966 ( NSW-135166). Morpeth, 1972, Johnson 7538 SW Oenothera indecora subsp. bonariensis reported in literature from outside of South America: SOUTH AFRICA: Jacot- -Guillarmod (1971); Ross (1972: 262). GERMANY: Léveillé & RANCE: Fournier (193 : 598, as O. argentinae ). Hybrids between O. indecora subsp. bonariensis and O. stricta subsp. stricta occur in Austr: alia € Pa ga NEW SOUTH WALES: Warialda, Lanagan in 1950 (NSW-135159). Casino mn Glenfield in 1950 (NSW. 135162). Aes ea Mair in 1953 (NSW-68289). Patagona Coans in 1954 (NSW-66109). Grafton, Flynn in 1954 (NSW-66107). Warialda, eni in 1956 (NSW-135158). Grafton, O'Grady in 1957 (NSW, RSA). Moree Distr., Mac- tier in 1962 (NSW-135155). Premer, s ^u 1962 (NSW-135157). Ashford, McNamara in 1963 (NSW-135160). Menangle Park, McBarron 9507 (NSW). Sackville Reach near Hawkesbury River, Walker in 1965 MN pe. Dee Why Laggon, Coveny in 1966 (NSW- 26 . Narrabeen Lake, Coveny in 1966 ( NSW-135165). Near Iluka, ENE of Maclean, 1969, Coveny 2178 (NSW, DUSS B PORTUGAL, ESTREMADURA: Me 1954, Rainha 2703 (COI, LISE), 1961, Rainha 4847 (LISE); Caldas da Rainho, 1960, 5 i 4495 (COI, LISE); Costa da Caparics, Matos in 1967 (COI). Despite its extensive range, O. indecora subsp. bonariensis is very uniform, in contrast, for example, to O. odorata with its seemingly endless variability. This might be related to the great ecological amplitude of O. odorata and the 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 523 Fi ; 166. NA punae (Bolivia, La Paz, Santarius 2013); Oenothera sect. Oeno- thera ate Munzi much more limited range of habitats in O. indecora subsp. bonariensis, which flourishes only in sandy places. At least a major portion of the chromosomal complex of O. indecora subsp. bonariensis is represented in the chromosomally heterozygous entities O. picen- sis subsp. bonariensis and O. parodiana of series Allochroa, as well as in O. brevi- petala, O. tucumanensis, and O. punae of series Clelandia. The chromosomal heterozygotes from Uruguay (Santarius 68, 72) have two complexes derived within O. indecora. The relatively short bracts of these plants suggest, however, that one of these complexes may be modified by genes from O. ravenii, probably transmitted to O. indecora via the small-flowered O. parodiana. 19c. Oenothera indecora subsp. boliviensis Dietrich, subsp. nov. Plantae maxime ad 20 cm altas, bene ramosae. Pubes ut in subsp. bonariensis. Folia caulina 2-3 cm longa, 2-3 mm lata; bractea 1.5-2 cm longa, 1.5-2 mm lata. Gemmae 2-3 mm longae, 1.5-2 mm crassae. Petala 2-3 mm longa. Capsula 1.5-2 cm longa. Semina ambito elliptica, 0.8-1 mm longa, 0.3-0.4 mm crassa. Cleistogama. Numerus gameticus chromosoma- ticus, n = 7; planta chromosomatice homozygotica. Plants attaining a height of only 20 cm, bushy and well branched. Pubes- cence as in subsp. bonariensis. Cauline leaves 2-3 cm long, 2-3 mm wide; bracts 1.5-2 cm long, 1.5-2 mm wide. Buds 2-3 mm long, 1.5-2 mm thick. Petals 2-3 mm long. Anthers 1.5-2 mm long. Filaments 1.5-2.5 mm long. Capsule 1.5-2 524 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 64 cm long. Seeds elliptic in outline, 0.8-1 mm long, 0.3-0.4 mm thick. Cleistoga- mous. Gametic chromosome number, n = 7 (7 bivalents* at meiotic metaphase I). Type: Cultivated in the Botanical Garden of Düsseldorf, Germany, 8 Aug. 1972. Source: Bolivia, near La Paz, A. Hecht 1964-29 (MO-2155715, holotype; CTES, DUSS, M, isotypes). Distribution (Fig. 233): Known only from the type locality. Specimen mme. from cultivated plants BoriviA. LA PAZ: La Paz, Hecht 1964- 99* (CTES, DUSS, M, MO). This subspecies seems clearly to be a derivative of subsp. bonariensis and is thus far known only from cultivation. It might very possibly be a chromosom- ally homozygous derivative of the complex heterozygote O. punae, which in turn has been derived by the combination of various species of series Renneria with O. indecora subsp. bonariensis. The range of subsp. bonariensis extends to high enough elevations in the province of Salta so that this combination is pos- sible. If this reasoning is correct, then O. indecora subsp. boliviensis provides an excellent demonstration of the way in which a genome can be altered when it is combined in a complex heterozygote. The F, generation derived by hybrid- izing O. indecora subsp. bonariensis with O. punae segregated, as might be expected, into two distinct classes of plants, one of which has the habit of O. punae, the other the more normal habit of O. indecora. This indicates that the normal habit of subsp. bonariensis is dominant over the nanism of subsp. boliviensis. The absence of O. indecora subsp. boliviensis as a wild plant might be explained on the hypothesis that its genome can exist at high elevations only in combination with those derived from series Renneria. It should however be sought in the vicinity of La Paz, Bolivia. 90. Oenothera affinis Cambess. in St.-Hilaire, Fl. Bras. Mer. 2: 269. 1829.— Fics. 58-59, 133, 180-181, 218. O. otaq Spach, Nouv. Ann. Mus. Hist. Nat. 4: 343. 1835. type: Chile, Prov. Valparaiso, , Bertero (P, holotype, F and GH photographs; G, isotype); Ann. Sci. Nat. Bot., Sér. 273. 1835. O. chilensis Fischer & Meyer, Ind. Sem. Hort. Bot. Petrop. 45. 1835. rEcrorvrE: Cultivated e Botanical Garden at Leningrad from seeds of the Botanical Garden at Paris, 1835 (LE). O. macrosiphon Lehm. ex Otto, Hamburger Garten- Blumenzeitung 14: 439. 1858. Based on nts cultivated at Hamburg, the seeds received from Darmstadt as O. villosa; authentic 1 not seen. A contemporary specimen bearing this name, probably cultivated in Ber in, was in the K. Koch herbarium in Berlin until it was destroyed in World War II; a photograph and fragments are at POM. O. mollis ssima L. var. grandiflora Micheli in Mart., Fl. Bras. 13(2): 178, tab. 38. 1875, pro 8 "rà 5 H. Lév. race propinqua (Spach) H. Lév. var. 5 (Spach) H. v., Monogr. Onoth. 364. 1909; Bull. Acad. Int. Géogr. Bot. 19: 9. Raimannia berteriana (Spach) e & Riley, p Misc. pa 1921: du 1921. 5 0 diplotricha Domin, Bibl. 22 (89); 1928. PE: 5 South-Queens- land, mixed forests at Logan E on sand, 2n K. Danis HI. Not located. O. mollissima sensu Munz, Physis 11: 282. 1933, pro parte; Amer. J. Bot grs 659. 1935, pro parte; Revista Univ. Chile 22. 1: 266. 1937, pro parte; Comun. Bot. Mus. Hist. Nat. Montevideo 1(10): 29. 1943, pro parte. H 1977] DIETRICH—SOUTH AMERICAN OENOTHERA Ficure 167. Oenothera laciniata subsp. pubescens (Peru, Junin, Santarius 2190). O. mollissima “El Cuadrado” Cleland, Jap. J. Genet. 43: 332. 1968. Oenothera affinis “longiflora Erlangen” Cleland, Jap. J. Genet. 43: 332. 1968. Erect annual herb, not forming a rosette, unbranched or + well branched throughout, 4-15 dm tall. Entire plant covered with soft hairs, densely to sparsely long-villous, the hairs erect, densely short-villous, and densely glandu- 525 526 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 lar-pubescent. Cauline leaves cultrate to narrowly lanceolate, acute, acute to rounded at the base, sessile, 5-15 cm long, 0.5-1.5 cm wide; bracts cultrate to narrowly lanceolate, acute, rounded to truncate at the base, sessile, longer than the capsules they subtend, (3-)4-9 cm long, 0.5-1.2 cm wide; leaves plane or weakly to evidently undulate at the margins, sparsely serrate with blunt teeth. Inflorescence branched. Floral tube 8-11(-13) cm long. Buds narrowly lan- ceolate to lanceolate in outline, green or yellowish green, often flushed with red, 2-3.5 cm long, 6-9 mm thick; apices of the sepals erect or divergent, 1.5-4 mm long. Petals very broadly obovate, (1.5-)2-4 cm long. Anthers 10-14 mm long. Filaments 15-20 mm long. Style long or short, the stigma held above the anthers at anthesis or surrounded by them, 9-13.5 cm long. Stigma lobes 5-10 mm long. Ovary 1.3-2 cm long. Capsule (2-)2.5-4(-5) cm long, 3-4 mm thick, thicker in the upper third, and with the 4 valves clearly separated at the apex. Seeds elliptic in outline, 1.5-2 mm long, 0.5-0.6 mm thick. Self-compatible; outcross- ing or self-pollinating. Gametic chromosome number, n = 7 (7 bivalents*, ring of 14** or intermediate configurations at meiotic metaphase I). Flowering time: Brazil, Uruguay and eastern Argentina, September-May; central Argentina, October-April; Bolivia, Chile and northern Argentina, November-April. Type: Brazil, Rio Grande do Sul, margins of woods near city of Rio Pardo, April (1816-1821), Auguste de St.-Hilaire 2791-12 (P, holotype, F and GH photographs). Distribution (Figs. 234, 244): In Brazil from Rio de Janeiro through Minas Geraís to Rio Grande do Sul. Apparently absent in the Atlantic departments of Uruguay, but frequent in the western, central, and southern departments of Ártigas, Salto, Paysandu, Río Negro, Soriano, Colonia, Flores, San José, Tacua- rembo, Durazno, Florida, Canelones, and Montevideo. From northern Argen- tina the species extends to Tarija in Bolivia. It occurs throughout northern Argentina, where it ascends to an elevation of 2,500 m along the river valleys of the Andes, and in central and eastern Argentina. It occurs in the following provinces of Argentina: Jujuy, Salta, Tucumán, Catamarca, La Rioja, San Juan, Mendoza, Chaco, Formosa, Santiago del Estero, Santa Fé, Córdoba, San Luis, La Pampa, Misiones, Corrientes, Entre Ríos, and Buenos Aires. West of the Andes, the species occurs in Chile from the province of Atacama to Valparaíso. Specimens examined from cultivated plants URUGUAY. MONTEVIDEO: Garden of the faced de Agronómia in Montevideo, Santarius 193* (ring 6, 4 bivalents), 194-196, 197*, 198-204 (DUSS; 193, 197 also CTES, M, MO). Toledo, Hecht 1964-95** (CTES, DUSS, M, MO). Botanical Garden 5 the Facultad de Agronómia in Montevideo, Rosengurtt B11256 (ring 6, 4 bivalents) (DUSS). FLORIDA: At the railroad about 1 km NE and 3 km SW of Mansavillagra, Santarius 215, 217, 218 (ring 12, l bivalent), a 222* (ring of 8, 3 bivalents), re , 224, 225*, 226, 228*, 229, 230 (DUSS; 218, 222, 225, 228 also 791505 M, MO). 3 km of Arrayän at Ruta 7 near the Rio Mansavi- llagra, Santarius 2n (D USS). Florida, Hecht 1964-1** (DUSS, M, MO). RGENTINA. BUENOS AIRES: Garden of the Botanical Institution of the Facultad de Agro- nómia in Buenos Aires, Santarius 259, 261 (5088) Old gravel mine E of the road from Berisso to Los Talas, about 1 km SE of Villa Zula near La Plata, Santarius 300*, 301*, 302, 304, 308-312, 314*, 315-317, 319, 321, 324-326, 328 (DUSS; 300 also CTES, M; 300, 310 also MO). Punta Lara near La Plata, Gopel in 1961 (DUSS); iui 1964- (DU SS). Bo- tanical Garden of Buenos Aires, Diers 19* (DUSS, M). Villa or in Buenos Aires, Hecht 1964-125* (CTES, DUSS, M, MO). ENTRE níos: Gualegu ar Parque ae Burkart 23060 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 527 (DUSS). Isla Almiron Chico near Concepcion del Uruguay, Burkart 23062 ( DUSS). eT FE: Santa Fé, Cleland 1965-426* (DUSS). cónpoBA: Córdoba, Hecht 1964-90 (DUSS). Cuadrado, Cleland 1967-417** (DUSS). sarra: Dep. Cafayate, Tolombón near the be of acumen. 1,600 m, Hunziker 21104 (CTES, DUSS, MO). jujuy: In the bed of Rio Xibi S of San CREMA de Jujuy, 1,250 m, Santarius 1841**, 1842, 1843 (DUSS; 1841 also MO). In the bed of Río Grande N of San Salvador de Jujuy, 1,250 m, We 1849** 1850* (DUSS; 1850 also CTES, M, MO). Ro: 5 to Pucara, 1 km S of Tilca 2,450 m, 1 1851 * 1852“ (DUSS; 1851 also CTES, MO). Damp id NE of Tilcara, about 150 m of the cemetery, 2,500 m, Santarius IE 1854, 1855, 1857, 1860, 1863, 1867, 1869*, m 1872, 1877, 1878, 1881, 1882. 1884 (DUSS; 1869 also CTES, M; 1869, 1882 also MO). rucu- MÁN: Stony places, Río Salí, near the bridge of Ruta 9, E of Sab Miguel de Tucuman, 420 m, Santarius 1672**, 1674, 1675* (DUSS; 1672, 1675 also M, MO). Bed of Rio Lules, N and NW of Lules, 410 m, Santarius 1679*, 1680 (ring of 6, ring of 4, 2 is 77 1682, p 1687, 1693, 1695, 1697, 1699, 1702, 1709, 1711* yum of 4, 5 bivalents) (DUSS; 1711 also CTES, M; 1680, 1697, 1711 also MO). Bed of Río Seco near bridge at Ruta 38, 450 m, Santarius 1712 (ring of 8, 3 bivalents) (DUSS). Bed of Rio Gastona near pe bridge at Ruta 450 m, Santarius 1713 (ring of 6, 4 bivalents (DUSS, MO). Bed of Rio Angostura, pior Tafi del Valle t to 5 km N, 2,000-2,250 m, ce 1714*, k 1722, "1724. 1727, (DUSS; 1714 also CTES, M; 1714, 1727 also MO). Bed of Rio Angostura at km 24.5 of i 307 near Tafi del Valle, 800 m, Santarius vis. (ring of 8, 3 bivalents), 1789-1791, 1794, 1799, 1800 (DUSS; 1788, 1799 pw MO). At km 16.5-17, 570 m, Santarius 1801 (ring of 8, 3 bivalents; ring of 8, ring of 4, 1 bivalent) (DUSS, MO). At km 11 in the bed of Río Toto- rilla, 430 m, Santarius 1802 (ring of 4, 5 e 1803, 1804 ( DUSS; 1802 also MO). Cla- villo near Tucumán, Cleland 1967-425* 5 USS, M, MO). LA Roja: Costadero to Mina, Hecht 1964-1 20* (CT ES, DUSS, M, MO, HILE. 5 Dry place at a road ina PA near Pisco del Elqui, Stubbe in 1961 (ring of 12, 1 biva ULTIVATED. 0. “longiflora” from the Botanical Garden in Erlangen, Germany, received 1962* (CTES, DUSS, M, MO). Representative specimens examined: BRAZIL. MINAS GERAIS: Caldas, pur 2799 (POM, SP). Between Carandaí and Crespo, Duarte 524 (BR). Morro de Pd d near Ouro Preto, Emygdis 2881 (R). RIO DE JANEIRO: Villa Theresa, Glaziou 8344 (G, K, LY, P). são PAULO: Pinheiros, Usteri 11896 (SP). Sao Paulo, Hoehne 11670 (POM). acies Galvão in 1884 (R). Belémsinho, Porto 11670 (SP). PARANÁ: Oure Fine near Bocayüva do Sul, p on 963 (HB). Rio Ja ngada near Palmas, Hatschbach 3493 (RSA). Serro = Rambo 9 (SP). Rio Cavernoso near Guarapuava, Pereira 7696 (HB). Rio Perdido near ns d do Sul, Hatschbach 19855 (UC). sANTA CATARINA: Garopaba, Klein & Bresolin 8850 (HBR). Valley of Rio Pelotas, Pabst & Pereira 6224 (HB). Lajes, Morro do Pinheiro Seco, 1,000 m, Reitz 6601 (HBR). Itapiranga, 1 71 m, Klein 5200 (HBR, RSA). AM of Rio Uruguai near gm Smith & Reitz 9728 (HBR, R, RSA, US). Ubatuba, Hans 311 (R). RIO GRANDE DO SUL: Banks of Rio Ibira Eo near Alegrete, Palacios & Cuezzo rm (LIL). Rio Pardo, m s 16 (B). Serro Largo, Sehnem 3549 (SI). Between S. Angelo and Guarani das Missóes, Santos 2727 (R). RUGUAY. ARTIGAS: Arroyo Tres Cruces Grande, Praderi 2540 (LIL). sALro: San An- tonio, Osten 5423 (SI). PAvsANDU: M. Cassioni, Puerto 8303 (MVFA). soriano: Fray Ben- tos, Fruchard in S (P, RSA, US). Sta. Elena, Rosengurtt PE-4386 (GH, LIL, MO, SP). COLONIA: Colonia Valdense, Dubugnon 77 (G). TACUAREMBO: Valle Edén, Arrillaga 1733 A). DURAZNO: Bebween R io Negro and Est. km 329, Ziliani vip (MVFA). FLORIDA: Mansavillagra, Rosengurtt B791 (GH, POM). rronEks: Rio Yi, Rosengurtt B564 (POM). sAN JosÉ: Barra Santa Lucia, Osten 4551 (G). Arazati, Rosengurtt 1696 (POM). CANELONEs: Arroyo de Sauce, Bartlett 20963 (GH, MICH, NY, P, UC, US). MONTEVIDEO: Sta. Lucía, pee in 1972 (P). Colón, Osten 3714 (G). Montevideo, Isabelle in 1838 (W). Av. Alto Paraná, Montes 9929 (LIL). OLIVIA. TARIJA: Tarija, Fries 1125 (S); Le. 41 (LIL); 2,000 m, Fiebrig 2435 (BM, G, K). Padcaya, 2, 100 m, Fiebrig 2545 ( BM, G ARGENTINA. JUJUY: Yala, 1,600 m, Arnow 3747 (MO ). Tilcara, ca. 2,400 m, Balls 5955b (F, K, UC, US); Meyer in 1940 (F, GH, LIL-33737, NY, SI). Quebrada de la Huerta near . eee 2,700 m, Cabrera 12041 (BAB). El Fuerte near Sta. Baebara, Cabrera 17273 BA . SALTA: La Candelaria, 1,200 m, Schreiter 9436 (GH, LIL). Pampa Grande, Eier 6906 (BAB). El Morenillo near Rosario de la Frontera, 690 m, Carbone 10088 (LIL). Victoria, Meyer 5040 (LIL, UC). Cachí, Garolera-Romero in 1947 (LIL, RSA). 528 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Tolombón near Cafayate, 1,600 m, Hunziker 21104 (CORD, MO); Hayward 002066 (LIL). TUCUMÁN: Rio Churqui, Lillo 7512 (LIL, US). Rio Sali near Tucumán, 600 m, Venturi 999 (GH, LIL, POM, SI, US). Rio Cochuna near Chicligasta, 915 m, Munz 15476 (GH, NY, POM, US). T Tafi del Valle, Araque & Barkley 19Ar161 (F, LIL). Cerro Munoz near Tafí, 2,500 m, Lillo 1241 (LIL). Rio Lules, 760 m, Mexia 04354 (GH, MO, UC). Rio Tipamayo near Trancas, Venturi 4218 (GH, LIL, NY, POM, US). S. Pedro de Colalao near Trancas, Krapovickas & Cristóbal 15339 (CTES). Cerro del Campo near Burroyacú, 2,000 m, Venturi 7748 (K). CATAMARCA: Andalgala, Jorgensen 1054 (BAB), 1055 (GH, LIL, MO, POM, SI); Rodriguez 344 (GH, LIL, S, UC). Rio Pomancillo, 5 29563 ( BAB). San Antonio near Padín, de Ance in 1957 (SI). El Portezuelo, Bartlet 19613 (MICH, ay Yacutula near Belén, Schickendandz 131 (CORD, GOET). Barranca i near Belén, 2,400 m, Schreiter 10248 (LIL). Near Capayan, Risso 556, 565, 626, 770 (LIL). Sta. Maria, 8 3,000 m, Peirano in 1933 (LIL). Sierra de Ambato, between Mutquin and Poman near Colana, 1,650 m, Hunziker et al. 18406 (CORD, MO). LA nioJa: Sierra Famatima, Alto Carrizal, ia nh 1044 (BAA). La Hoyada, 2,500 m, Kurtz 15041 (CORD, MO). Guanchin Viejo, Castellanos 28-284 (BA, POM). Santa Cruz, Toscani 32 (BAB). Campanas, 1,850 m, Rojas Paz in 1942 (GH, N85 970 Rodeo de Las Vacas, 3, 000-4, 000 m, Flossdorf 40 (S D. Dep. Capital, El Cantade 200 m 17 00 3939 (GU, LIL, UC). Sierra de Velazco, El Cantadero, 2, rH m, Hon" |. (CORD, SI). Yacuchi, Kurtz 15414 (MO). Punta de Agua near Gral. Sarmi- ento, 2,600 m, sind 2083 (CORD, POM). Los Molles near Lamadrid, 2,700 m, Pee vickas & Hunziker 5530 (BAB, CORD). san JUAN: Between Rodeo and Colanguil near Iglesias, rag ain ed (US). Dep. Calingasta, Las Lumbreras, 2,000 m, Spegazzini 656 (BAB). Río Calingasta near Calingasta, Zarchini 150 (CTES). Quebrada de Huachi near Jachal, 1,500 m, "piv 2993 (LP). Chimbas near Desamparados, Cuezzo 1266 (LIL, RSA). Valle Fertil, Hunzike er 16686 (CORD). Villa Krause near Pocitos, Cuezzo 2175 (LIL). La Bebida near Rivadavia, Cuezzo 1433 (LIL). MENpoza: Philippi 614a (SGO). Junin, Ruiz Leal 7033 (Leal, LIL). rermosa: Hicken (SI-4851). Pirané, Morel 638 (LIL). Uriburú at Rio Bermejo, Kermei 68660 (BAB). cHaco: Las Palmas, Jörgensen 2488 (GH, LIL, SI, US). Resistencia, Meyer 3212 ( LIL, POM). Fontana near Resistencia, Meyer 456 (LIL, SI). Colonia Benítez, Stuckert 19511 (CORD, G); Schulz 501 (POM). Fortín Anguilar, Hossens in 1917 (CORD). SANTIAGO CEL ESTERO: Dep. Rivadavia, between Selva and Palo Negro at Ruta 34, 100 m, Hunziker 17776 (CORD, MO). santa FE: Mocoví, Venturi 275 (SI); Qua- rin 1145 (CTES). Monje, Pedelaborde in 1940 (LIL, SI). At Ruta 9 across the Río Caraca- raná near lriondo, Hunziker 13702 (CORD, MO). Reconquista, Venturi 275 (LIL, POM). Las Colonias near Pilar, Terribile 474 (LIL). Aurelia near Castellanos, Terribile 490 (LIL). CÓRDOBA: Ban Río Primero near Ojo de Agua, Hunziker 11920 ( CORD, MO). Gral. Paz, Stuckert 3730 (CORD, G, MO). Sierra San Ignacio near Quintas, Stuckert 18586 (CORD, G). ique Los Molinos near Calamuchita, Krapovickas & Cristóbal 14714 (CTES). Río Tercero, Burkart 10369 (MO, SI). Near Santa María, Krapovickas 6514 (BAB, CORD, LIL). Capilla del Monte, Nicora in 1941 (SI-17600). Totora l, 5 106 (POM). La Carlota near Juárez Celman, Hossens 385 (CORD). Córdoba, ne tz 308 (CORD, GOET); Hieronymus in 1878 (FR); Lossen 128 (G, LE, M, Z). 3 Giardelli 845 (SI). Las Com- puertas at Rio Segundo, Sublis 1171 (CORD, M Siena Chica, between Pan de Azúcar and Villa Allende, 1 11932 (RSA). Cruz del Eje, Meyer 13042 (LIL). San Javier, Cas- tellanos 10734 (RSA); Fabris & Moreau 6791 (BAB, LP). Capilla del Monte, Hossens 427 CORD). lim Hieronymus 333 (P). Los Cocos near Punilla, De la Sota 3278 (LIL). SAN LUIS: Sierra de Comechingones, El Rincón, Hunziker 11852 (CORD, MO). Piedra Blanca near Merlo, 1,000 m, Digilio- Grassi 2066 (LIL, RSA). Trapiche, Gez 31-278 (POM). Que- brada de los Bueyes, Galander in 1882 (CORD). LA PAMPA: Between Chamaico and Casi- miro Gomez near Rancul, Cabrera & Sagastegui 19406 (LP). Misiones: Corpus near San Ignacio, Schwarz 3411 (LIL, S). Gob. Roca near San Ignacio, Schwarz 6413 (LIL). San Ignacio, Quiroga in 1913 (POM). San Pedro, Bertoni 2101 (LIL). Victoria at Ruta 12 near Iguazu, 170 m, Schmidt 2743 (LIL). Bonpland near Posadas, Lillieskéld (F, S). Garupá near Cainguas, Bertoni 4660 (LIL). Santa Ana, Rodriguez 109 (GH, LIL, SI, UC). 1 Ruta 105, Mroginski 430 ( CTES, MO). corrientes: Mercedes, Rodrigo 711 (LP, N Ist. La Pastoril,“ at Rio Paraná near Lavalle, Pedersen 3858 (BR, C, G, UC, US). Monte 1 9 Nicora 5748 (BAA, CTES). Paso de los Libres, Ibarrola 2002 (LIL, NY, S); Schinini 7698 (CTES, MO). Santo Tomé, Ibarrola 1233 (LIL, NY). Curuzú Cuatia, Spegazzini 154 (BAB). Bella Vista, Schinini 6565 ( CTES, MO). ENTRE Rios: Concepción del Uruguay, Lorentz 628 (BREM, COI, CORD, F, FR, G, HBC, K, L, LE, LY, M, UPS, W, Z), 12 (GOET). Arroyo 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 529 Martinez, Boelcke 907 (BAA). Paraná, Frommel & Lefebre in 1888 (P). Paracao near Parana, Burkart 23762 (MO). Calera Barquin near Colón, Pozzi 27-1572 (POM). La Paz, Burkart 21312 (SI). Federación, Meyer 11130 (LIL). e Burkart 22243 (MO, RSA). Isla Almiron Chico, Burkart 23062 (MO). Gualeguaychú, Burkart 23060 (MO). BUENOS AIRES: Delta, Río Carabelas near Tiburón, Scala in 1925 (NY). Est. Las Palmas near Zárate at Río Paraná, Boelcke 13166 (BAA, M O). Hudson near La Plata, Eyerdam & Beedle 23156 (G, GH, K, MO, UC). Pereyra near La Plata, Cabrera 7607 (LP). Palermo, Munz 15460 (NY, POM, US). Los Talas near Berisso, Boffa 171 (F, LIL, S). Atucha near Gral. Uriburu, Kra- povickas 3290 (BAB, LIL). San Isidro, Parodi 8565 (GH). Martín Garcia, e 5034 (SI). Otamendi, Fabris 4995 (LP). Pergamino, Boelcke 2215 (BAA). Punta Lara, Dawson 854 „ GH, LP, NY); Rodríguez 520 (S, SI). CHILE. ATACAMBA: Valley of Rio Transito, La Pampa near Vallenar, u 5860 (GH, K, POM). Higuerita near San Félix, 1,300 m, Ricardi 3867 (CONC). coquiMBo: Gay in 1838 (P). Rivadavia, 800 m, Wordeman 175 (BN {, F, G, GH, HBG, LIL, "MO, SI, UC, Z). Rapelcillo near Ovalle, 800 m, Jiles 1958 (CONC). Between Pauhuano and Elqui, Pfister 6498 ONC). Tunga near Mapel, Landbeck in 1962 (SGO). Aconcacua: Los Andes, 1,100 m, Behn 22802 (CONC); Scott Elliot 436 (BM); Philippi in 1885 (HBG, SGO). Pun Felipe, Claude-Joseph 2508 (US). Chupaja, Jelinek (W). Near Papudo, Zóllner 5168 (L). var- PARAÍSO: Petre in 1818 (S), in 1827 (W). Laguna Verde, 100-150 m, Eyerdam et al. 10040 (F, NY, UC, US). Concón, Poeppig 121 (BM). Limache, AJ.H. in 1927 (M). Quillota, Maximowitsch 133 (LE). sANTIAGo: Philippi S86 (P, LE). Salto de Conchali, Philippi 614b (SGO); Marques in 1878 (HBG). Cordillera de Santiago, cba in 1856-57 (FR, G, W). Cerro de Renca, Gusinde 643 (W). ISLA JUAN FERNANDEZ; Cumberland bay, valley al Lord Anson, Bock in 1931 (SGO). Province unknown: Aeuleo, Bertero 464, 1185 pro parte ( BM, W). Farillar, Volckmann 28 (SGO). Specimens from outside of South America ace ScoTLAND. Selkirk, 1965, Webster 10169 (BM, SPAIN. i station of San Sebastian, Cuülon in 1877 (MPU). Ona, Arraiano in 1881 Lh ORTUGAL. ESTREMADURA: Sandy places near Lagoa Obidos, Daveau in 1882 (BM, COI, LISU). Samouco, 1882, Coutinho 1273 (LISU). Barreiro, da Cuba in 1888 (LISE, LISU). Trafaria, Daceau 3093 ge LISU); Jonge in 1912 (BAS). Moita, da Cunha in 1889, in 1891 (LISE), in 1890 (LISU). Brejo do Cobre, Santoz in 1905 (LISU). Caldas da Rainha, in 1890 (COI); 1938, 8 A 14098 (LISE). Praia das Macás near Sintra, 1950, Rainha 1974 (LISE, UPS). S. Marinho do Porto, 1961, Rainha 5060 (LISE). ALGARcE: Faro, Gui- maraes in 1880, in 1882 (COI). Tavira, Daneau in 1890 (LISU). PAKISTAN. West Himalaya, Hazara, Duthie in 1899 (K); 1963, Nasir & Siddiqi 1821 (RAW). Inna. Punjab, Abbotabad, 1922, Drummond 20096 (K). Punjab, Manikeru village in valley ieee near Kulu, 1934, Parkinson 3923 (K). Prov. Madhya Pradesh, Balakot, 1959, Jafri & Ali 326 ) TRALIA. QUEENSLAND: Currumbiu Beach S of Brisbane, 3 in 1961 (K). More- ton near Southport, 1936, Pedle 79 (K). Bubya Mountains, 1944, Clemens 43807 (LIL). South Pine River near Moreton, 1964, Henderson H97 (K). NEW SOUTH WALEs: Bellbrook, Lengoth in 1891 (MEL-58126). Conjola, McHeron in 1899 (NSW-135171). Rockdale, Camp- field in 1902 ( NSW-135174). Bega, Tielkens in 1906 (NSW-135172). Tweed Heads, Cheel in 1916 (NSW-135176). Sussex, Maiden in 1917 (NSW-135170). Urunga, 1917, Lawrence 8132-17 (NSW). oe Gillings in 1920 (NSW-135167). Wondabyne to Woy Woy, Blakely in 1922 (NSW-135173). Bomaderry, Rodway in 1936 (NSW-135175). Moura, Rod- way in 1936 (K, NSW- 21 Gearge's Creek, upper Macleay River, Danis in 1941 (NSW- 135180). Grafton, O'Grady in 1951 (NSW-135178). Kyogle Distr., Vane in 1958 (NSW- 135177). At Manning River near Wingham, 1964, Salasoo 2820 (NSW). Merriwa, 1970, Raven et al. 25866 (K, MO, NSW). Bylong, ca. 30 mi NE Mudgee, 800 m, 1970, Raven et al. 25869 (MO, NSW, PERTH). N Milton on Princess Highway, 1970, Raven et al. 25894 (K, MO, NSW). Brogo, 12 km N Bega, 1 Briggs 3970 (K). 1 mi W Tabulam, n Salasoo 4609 (NSW). SOUTH AUSTRALIA: Encounter Bay near Port Elliot, 1895, H. 392 (MEL). WEST AUSTRALIA: Bunbury, Wickens in 1910 (BM). 25 mi E of Albany, Elder in e (NSW). Mi Lawley, Lyon in 1966 (PERTH). Hawau. Parker Ranch, 1929, Carter & Brown 182 (K). Mauna Kea, near pit where David Douglas was murdered, 1949, Degener et al. 20344 (K, MO). 530 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 SOUTH AFRICA. CAPE: At P.P. Rust, 1905, Rogers 4117 (PRE). Paarl, 1916, Smith 2667 (K), 1926, 2667 (K, PRE). Roadside near Faure, Distr. Stellenbosch, 1968, Parker 4390 (K). TRANSVAAL: Nylstroom, 1902, de Jongh 6529 (PRE). Distr. Pretoria, Scheerport, 1906, Leendertz 8295 (PRE). Rutzenburg, 1910, Leendertz 9568 (PRE). Transvaal, Davy 1248 (PRE). NATAL: Distr. Durban, Isipingo, 1961, Ward 3764 (PRE). Distr. Umzinto, Shelley Beach, 1967, Strey 7281 (K, M, MO, PRE). LesorHo: Morija, 1918, Dieterlen 1352 (P, PRE). Old specimens from plants cl ultivated in gardens: Munich, Germany, Herb. Zuccarini in 1835 (M; as O. undulata). Paris, in 1835 (P; as O. misit jm du Luxembourg, in 1837 (K; as O. chilensis). Vienna, Austria, in 1849, W as O. Oe notera affinis reported in literature from outside of South America: PORTUGAL: Raven (196 he ee A: Ross (1972: , AUSTRALIA: Bailey (1900: ET om 215) (as O. rc d Black (1909: 63; 1926: 427 (as O. longiflora); 1952: 638); Beadle et al. (1962: 173). More or less clavate capsules with the valves d separate at the apex, a long floral tube, and a dense vestiture of soft pubescence are the most obvious characteristics of O. affinis. In comparison with O. odorata, the variation is lim- ited. Relatively leafy plants with broad leaves occur throughout the range, espe- cially in the east, but less densely leafy ones in which the leaves are relatively narrow occur only in the northern part of the area of distribution—Bolivia and the provinces of Argentina along the eastern flanks of the Andes. Just as in O. odorata, there are included in O. affinis complex heterozygotes which are indistinguishable from the homozygotes of the same species. Both sorts of plants occur in the same populations in O. affinis, together with all pos- sible intermediates between 7 pairs and a ring of 14. For example, plants from 2 vicinity of Tucumän had the following configurations: Santarius 1672, ring 4; 1675, 7 pairs; 1679, 7 pairs; 1680, ring of 6, ring of 4, 2 pairs; 1711, 7 pairs, 8 with ring of 4, 5 pairs; 1712, ring of 8, 3 pairs; 1713, ring of 6, 4 pairs; 1714, 7 pairs; 1788, 7 pairs, another with ring of 4, 5 pairs; 1801, ring of 8, 3 pairs; ring of 8, ring of 4, 1 pair; 1802, ring of 4, 5 pairs; Cleland 425, 7 pairs. See the general remarks in the introduction on p. 437. 21. Oenothera mollissima L., Sp. Pl. 346. 1753.—Fics. 60-62, 134, 182, 219. Onagra mollissima (L.) Moench, Meth. Pl. 1: 675. 1794. Oenothera mollissima L. var. villosa Sprengel, Pl. Min. Cog. Pug. Prim. 2: 60. 1815. TYPE: not seen O. holosericea Tausch, Flora 22: 558. 39. LECTOTYPE: Cultivated in botanical garden, 83 Tausch (PRC, POM 2 8 Munz, Amer. J. Bot. 22: 661. 1935. Onothera polymorpha H. Lév. race mollissima (L.) H. Lév., Monogr. Onoth. 365. 1909; Bull. cad. Int. Géogr. Bot. 19: 325. 1909. Oenothera mollissima L. var. genuina Hassler, Bull. Soc. Bot. Genéve, sér. 10 95 274. 1913. Raimannia mollissima (L.) Sprague & Riley, Bull. Misc. Infor. 1921: 201. Annual herb, not forming a rosette, the main stem erect or rising obliquely, unbranched or + well branched, with the side branches arising at right angles or + obliquely, 3-10 dm tall. Plants densely or very densely and softly long- and short-villous and densely glandular-pubescent. Lower cauline leaves very nar- rowly elliptic to narrowly elliptic, acute, narrowly cuneate to acute at the base, sessile, 4-7 cm long, 0.5-1.2 cm wide; upper cauline leaves and bracts narrowly oblong to lanceolate, acute, rounded to truncate at the base, sessile; bracts 2-4 cm long, 0.5-1 cm wide, longer than the capsules they subtend, as- 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 531 Ficures 168-176. Scanning electron micrographs of seeds of taxa of Oenothera sect. Oenothera subsect. Munzia.—168. O. peruana (Peru, Arequipa, Santarius 2106).—169. O versicolor (Peru, Junín, Santarius 2163).—170. O. lasiocarpa (Argentina, Tucumán, Diers in 1959).—171. O. santarii (Argentina, Mendoza, Santarius 1430).—172. O. longituba ( Argen- tina, Tucumán, Preis M 1736).—173. O. scabra (Bolivia, Cochabamba, 5 2003).— 174. M mendocinensis (Argentina, Buenos Aires, Santarius 415).—175. O. odorata ( Argen- tina, Rio Negro, Santarius 800).—176. O. ravenii subsp. ravenii (Brazil, Rio Grande do Sul, "apa in 1966). cending and overlapping towards the apex of the stem; leaves plane to strongly undulate along the margins, distantly serrate with blunt teeth. Inflorescence branched. Floral tube (1.5-)2-5 cm long. Buds oblong to lanceolate in outline, green or yellowish green, often red-striped at the junction of the sepals with the floral tube, 0.8-1.5 cm long, 3.5-6 mm thick; apices of the sepals erect, 1-2 mm long. Petals obovate to very broadly obovate, often broadly elliptic, 0.8-2 cm long. Anthers 4-8 mm long. Filaments 7-12 mm long. Style short, the anthers 532 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 shedding pollen directly on the stigma at anthesis, 2.5-6 cm long. Stigma lobes 3-5 mm long. Ovary 1-L3 cm long. Capsule 2.5-3.5 cm long, 3-4 mm thick, often slightly enlarged in upper third. Seeds elliptic in outline, 1.5-2 mm long, 0.7-0.8 mm thick. Self-pollinating complex heterozygote. Gametic chromosome number, n — 7 (ring of 14* or ring of 12 and 1 bivalent** at meiotic metaphase I). Flowering time: October-July. Lectotype: Grown in Cliffort’s Garden in Hartekamp, Netherlands, 1735- 1737, C. Linnaeus (BM; GH and POM photographs). Said to be from the fields of Buenos Aires, Argentina. Linnaeus's diagnosis in the Species Plantarum is taken directly from Viridiarum Cliffortianum (1737) so the species must be typified by material he had available at that time. Distribution (Fig. 232): Exclusively in the eastern portion of the range of series Allochroa. In Brazil it occurs in the two southernmost states, Santa Cata- rina and Rio Grande do Sul; in Uruguay, only in the coastal departments of Rocha, Maldonado, Montevideo, Florida, Canelones, San José, and Colonia; in Argentina, it is found in the provinces of Misiones, Corrientes, Entre Ríos, and Buenos Aires. Ta examined from cultivated plants: GU MALDONADO: Sandy place in 92 part of Piriapolis, Santarius 117*, 118-122, 123°, 124-199 (DUSS; 117, 123 nun FETES, M; 117, 118, 123, 128 also MO). S andy places in S part of Piriapolis, Santarius 130*, 131- 134, 135*, 136-138, 139*, 140-143, 144*, 145, 146, 148, 149, 151 (DUSS; 130, 144, 151 also CTES; 130, 144, also M; 130, 134, 139, 144, 151 also MO). Dunes E of Piriapolis, Santarius 157*, 158-173 (DUSS; 157 also M; 157, 166 also MO). Dunes in the W district La Pastora of Punta del Este, Santarius 174*, 175—185, 186*, 187-191, 192* (DUSS; 174, 186, 192 also CTES, M, MO). Dunes on the coast near Piriapolis, Krapovickas & Cristobal 11147* (DUSS, MO). wowrEvipEeo: Dunes D pers and Miramar, Santarius 2*, 3-8, 10, 13*, 14, 16, 19-22, 23*, 24*, 25, 26, 29-31 (DUSS; 2, 13, 23, 24 also CTES; 2, 3, 13, 23, 24 also M, MO). Dunes and pine forest in the nn F. D. Roosevelt, 3-4 km E of Carrasco, Santarius 32*, 33-35, 36*, 37, 38, 39*, 40, 41, 42*, 43*, 44— 47 (DUSS; 39, 42, 47 also CTES, M; 32, 36, 39, 42, 47 also M O). Carrasco, Hecht 1964- 61* (DUSS). Montevideo, Hecht 1964- 87* (DUSS). Toledo, Hecht 1964-88* (DUSS). CANE- LONES: Canelones, Hecht 1964-25* (DUSS, MO). coLoNIA: Sandy place, port of Juan L. Lacaze, Santarius 48* , 49, 50*, 51-53 (DUSS: 50 also CTES, M; 48, 50 also MO). At roads in W part of Juan L. ' Lacaze, Santarius 67* (DUSS). Sandy place in the N part of Juan L. Lacaze, Santarius 80, 92*, 93-98, 99* 100, 101* (DUSS; 99, 101 also CTES, M; 94, 99, 101 also MO). Dunes NW of Juan L. Lacaze, Santarius 102*, 103-108 (DUSS; 102 also MO). 57 A the Vries 5 km NE and SW of Mansavillagi ra, Santarius 214**, 221* (CTES, DUSS, MO; 214 : ARGENTINA. CORRIENTES: At Ruta 40 8 km N of Santo Tomé, Krapovickas d» Cristóbal 16406* (Doss, MO). BUENOS AIRES: Garden of the Botanical Institution of the Faculdad de — in Buenos Aires, Santarius 256*, 263, 264, 265*, 266, 267 (DUSS; 256, 265 also CTES, M; 256, 264, 265 also MO). Botanical Garden of Buen os Aires, Gópel in 1961* ( CTES, DUSS, M, MO). Villa Gesell in General Madariaga, Burkart in 1962* ( CTES, DUSS, M, MO). ATED: From the main school garden in Frankfurt, Germany, received 1959*, source unknown (CTES, s M, MO). From the Botanical Garden in Erlangen, received 1960* TES, DUSS, Additional Lui. examined: RAZIL. SANTA CATARINA: Campo de Massiambu near Palhoça, 5 m, Reitz 4880 (HBR), Reitz & Klein 1222 (HBR). RIO GRANDE po SUL: Fazenda Bernardo Velho near Bom Jesus, 1,000 m, Rambo 34863 (5): between Capáo da Canoa and Osório, Nelson in 1970 (BR); Torres, Rambo 56194, 54791 (HBR), Burkart 25105 (SI); city of Rio Grande, Malme 244, 244a, 270 (S); S. Leopoldo, Rambo 1283 (LIL); Ilha dos Marinheiros, Schwacke 2599 (R); Praia de Tramandahy, Vidal in 1913 (R 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 533 Uruguay. COLONIA: N.N. 28 (BAA); Riachuelo, Cabrera 3340 (NY); Arroyo de Pintos near Puerto Platero, Bartlett 20771 (GH, MICH NY, UC); Playa Fomento, Puerto 1041 MVFA). SAN josÉ: Barra de Sta. Lucia, Herter 168e (B); Eifler, Herter 168g (US, Z); Playa Pascual, Arrillaga 742 (MVFA). CANELONES: Toledo, Herter 168f (B, GOET, L, SP, ; Balneario San Luis, Zorrón 1718 (MVFA, P); Camelon Chico, Berro 5635 (MVFA); Atlan- tida, Osten 21655 (BREM, F, GOET), Barattini in 1940 (MO): Ruta Interbalneario at km 275, Lema 6750 (MVFA); Arroyo Sarandi, Izaguirre 149 (MVFA); El Pinar, Arrillaga 435 (MVFA); Las Piedras, Fruchard in 1869 (P); dunes near Floreta, Steer in 1923 (HBG). MONTEVIDEO: Rosa- Mato 1509 (LIL), Isabelle in 1838 (W), Chabataroff in 1939 (GH), Gibert 86b, 340, 342 (K), Gibert 1175 (W), Gaudichaud in 1839 (G); Carrasco, Munz 15441 (POM), 15449 (GH, POM), Felippone 2078 (SI), Kuhlmann in 1948 (BR); Buceo, Fruchard in 1875 (P); Playas Blancas, Fruchard in 1874 (P); nud . in 1876 (NY, RSA, S, SI, US), Herter 168a (LE, M, Z), 168b (G), 76266 (S); Cerro, Herter 168 ( LIL, MO, SI, UC); Punta Gorda, Rosengurtt B4331 (LIL, MO, MVFA, SP, US); Plata, Courbon in 1856 (RB); Malvin, Felippone 2325 (SI), Herter 168d E HBG, NY). ROCHA: La Paloma, Descole 174 (LIL); Santa Teresa, Léon 338 (BAA); La Ferera, Rosengurtt 9944a, 9944b (MVFA), Hossens 18 (CORD); Cabo Polonio, Hossens 113 (CORD). Mo- NADO: Berro 3667 (MVFA), Lourteig 168 (LIL), Costa 7199 (MVFA); Punta del Este, Descole 74 (F, GH, LIL, NY); dunes near Solis, Osten 22389 (S); Cerro Ingles, Osten 5300 (CORD, SI); Cerro San Antonio near Piriapolis, Pabst 5496 (HB); San Rafael, Descole 27 LIL); Punta Ballenas, Krapovickas & Cristóbal 12691 (CTES). Department unknown: Road As 5 Felippone 2642 (SI). I : Santa Ana near Candelaria, Montes 1517 (RSA); road to La Plan- "ue near "e Be s] Scala 258 (LIL); El Dor: ado near Iguazü, Schwindt 2231 (LIL); Tapiorny near Cainguas, 215 m, Schwinds 813 (LIL). ENTRE nios: Concepción del Uruguay, Lorentz 35 A rues BUENOS AIRES: Andersson in 1852 (S), Rohl 4497 (W); Villa Gesell near Gral. Madariaga, Boelcke 54 (BAA, MO), Burkart 22376 55 dunes near Pinamar, E 10099 (LP), 10674 ( LIL, LP); Dos 8 Sur, Molfino 785 ); Isla Paulino, Cabrera F); Punta Lara, Molfino 247 (POM), Pujales 54 (LIL); ne de Oro near Colón, 1 1100 (LP); isle of Martin Garcia, Moreau in 1933 (RSA); N. N. in 1949 (SI 26721), A: 35 (LIL); Necochea, Rodriguez 848 (LIL, S); Jauregui near Luján, Burkart 18493 ; Juancho near Gral. Madaxiaes. Pastore in 1936 (SI 26720), Cabrera 2715 (LP, NY); E Clemente, Cabrera 4262, 4271 (LP), Cabrera 4921 (NY), Krapovickas 132, 2874 (RSA); Lourteig 465 (GH, LIL), Krapovickas 1924 (LIL), Vervoorst 5208 BAB); between San Clemente del Tuyú and sy de Ajó, Tortosa & Medán 11058 (BAA); Samborobon near Gral. Lavalle, vndas 31-1630 (POM); Mar Chiquita, Lanfranchi 1822 (SI). ns from outside of South America (naturalized ): [ones STATES. New Jersey, Camden, with ballast, Martindale in 1866 {CH}, INDIA. Himalaya, cultivated at Almorah, Duthie in 1900 (K), Strachey & W interbottom = > 8 = — NEPAL. Cultivated (K). MET RALIA. Moers De Noosa Heads, Johnson “a 1951 (NSW 135152). NEW souTH VALES of Raymond Terrace, Coans in 1961 ( NSW 65373); NR Bay, Lithgow in 1965 (NSW Du iet Highway at crossing of Mirae River, 1 mi S of Albion Park, 0, Raven et al. 25887, 25896 (MO, NSW); Centennial Park, pe =i Johnson in 1971 135154). 97 (NSW Old specimens from plants cultivated in gardens: Erlangen, Germany, Herb. Schreber in 1779 (M). Paris, Herb. Cambessédes in 1781 (MPU). Vienna, Austria, in 1806 (W-14035). Schónbrunn at Vienna, in 1814 (W). Karls- ruhe, Germany, in 1817 (W). Dresden, Germany, in 1822 (W; as O. villosa Thbg.). Munich, Germany, Herb. Zuccarini in 1832 (M). Munich, in 1842 (M). enothera mollissima reported in literature froin outside ito miui are Pittier et al. (1947: 258; no material seen! ); LT "(1970 3 ea fig. 77; the illustration which is said to be O. mollissima, represents O. tetraptera, Cav., thera sect. Hartmannia.) Oenothera mollissima has two chromosomal complexes derived from O. affi- nis but is so different from that species in its small flowers and distinctive habit that it is best regarded as a distinct entity ( Tandon & Hecht, 1955, 1956). Hy- brids between O. mollissima and O. affinis or other chromosomally homozygous 534 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 species yield progenies in which the two classes of plants differ from one another only trivially, as shown by Haustein (1952). In some strains of O. mollissima, both complexes give rise to small-flowered progenies in various combinations; in others, one has genes for a flower size approximating that of O. affinis. The small-flowered trait is dominant over the large-flowered one. With O. mollissima 1 have included complex heterozygotes (for example, Santarius 214, 221) which are, on the basis of their broader leaves and longer floral tube, intermediate to O. affinis. Whether such populations have originated y hybridization between the two species and then become stabilized, or actu- ally represent an intermediate stage in the evolution of complex heterozygosity, is not known. Growing in the same populations as typical O. mollissima are often found forms which have apparently originated by introgression with the densely pubes- cent form of O. odorata which occurs in the Province of Buenos Aires. e introgression manifests itself in the lower stature, denser pubescence, undulate leaf margins, and shorter floral tube of these plants. It is not desirable to segre- gate them taxonomically, however, since they always occur mixed with other plants typical of O. mollissima and are best regarded as constituting a part of that species. The very high degree of self-pollination in this species leads to the stabilization of numerous distinctive lines both in the field and in the experi- mental garden, a veritable feast for the geneticist. Despite this, occasional out- crossing does occur and leads to recombination within the populations, as well as rare interspecific hybridization, which seems to have been of importance in enriching the variability of the species. 22. Oenothera rivadaviae Dietrich, sp. nov.—Fics. 10, 63-64, 135. Herba ut videtur annua, erecta vel prostrata, rosulata, simplex vel ramis prostratis vel ascendentibus e rosula, caulis principalis 3-7 dm longus. Plantae sparse strigulosae, dense ve sparse villosae, denseque vel sparse glanduloso-pubescentae. Folia rosulae linearia vel angus- tissime elliptica, acuta, lamina in petiolum gradatim decrescens, 10-15 c m longa, 0.4-0.7 cm ata; folia Vs linearia e anguste J r acuta, basi ass cuneata vel acuta, sessilia, 4—10 cm long, 0.5-0.8 cm lata; bractea anguste lanceolat ata vel 5 acuta, basi truncata vel 5 quam capsulam subte antam oe vel brevioria, 2-5 cm longa, 0.5- 1 cm lata; folia ad margines exigue ad valde undulata, plerumque irregula riter 1 ser- rata, Intloresceutis ramosa. Tubus floralis 1-15 c 1 longus. Gemmae ambito oblongae ve lanceolatae, virides, saepe rubellae, 05- cm longae, 2.5-4 mm crassae; apices e divergentes vel erecti, 2-3 mm longi. Petala une obovata, 0.6-1.5 cm longa, canarina vel citrina. Stylus brevis, stigmate sub id 'si antheris circumdatus. Ovarium 1.5-2 cm longum. Capsula 3.5-6 cm longa, 2-3 mm lata. Semina ambito anguste elliptica vel elliptica, 1.4—1.8 mm longa, 0.5-0.8 mm crassa. Numerus gameticus chromosomicus, n — 7; planta chromoso- matice heterozygotica complexa — Plants probably annual, erect or prostrate, forming a rosette, unbranched or with prostrate or ascending side branches from the rosette, the main stem 3-7 dm long. Plants sparsely strigillose, densely or sparsely villous, and densely or sparsely glandular-pubescent. Rosette leaves linear to very narrowly elliptic, acute, gradually narrowed to the petiole, 10-15 cm long, 0.4-0.7 cm wide; cauline leaves linear to narrowly oblanceolate, acute, narrowly cuneate to acute at the base, sessile, 4-10 cm long, 0.5-0.8 cm wide; bracts narrowly lanceolate to 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 535 Q Figures 177-185. Scanning electron micrographs of seeds of taxa of Oenothera sect. Oenothera subsect. Munzia (continued).—177. O. longiflora subsp. grandiflora yr apes Corrientes, Krapovickas d» Cristóbal 11293) 178. O. indecora subsp. indecora (Urugua Maldonado. Santarius ms 179. O. indecora subsp. bonariensis ( Argentina, Buenos ree Santarius 275 ).—180. affinis (Uruguay, Montevideo, Santarius 193).—181. O. affinis (Santarius 193).—182. P mollissima (Uruguay, Montevideo, Santarius 2),—183. O. elon- gata (Bolivia, La Paz, Santarius 2026).—184. O. punae (Argentina, Tucumán, Santarius 1742). —]185. O. verrucosa (Peru, Arequipa, Santarius 2068). lanceolate, acute, truncate to subcordate at the base, longer or shorter than the capsule they subtend, 2-5 cm long, 0.5-1 cm wide; leaves slightly to markedly undulate at the margins, mostly irregularly serrate with blunt teeth. Inflores- cence branched. Floral tube 1-1.5 em long. Buds oblong to lanceolate in out- line, green, often flushed ~ red, 0.5-1 cm long, 2.5-4 mm thick; apices of the sepals divergent or erect, 2-3 mm long. Petals very broadly obovate, 0.6-1.5 cm 536 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 long, rich yellow to pale yellow. Anthers 4-6 mm long. Filaments 6-8 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 1.8-3 cm long. Stigma lobes 4-5 mm long. Ovary 1.5-2 cm long. Capsule 3.5-6 cm long, 2-3 mm thick. Seeds narrowly elliptic to elliptic in outline, 1.4-1.8 mm long, 0.5-0.8 mm thick. Self-pollinating complex heterozygote. Gametic chro- mosome number, n — 7 (ring of 14* at meiotic metaphase I). Flowering time: November oonan e: Grown from seeds and cultivated in the Botanical Garden of Düssel- ine Germany, 18 July 1969. Source: Argentina, Prov. Chubut, gravelly and sandy places in Villa Balneario Rada Tilly, ca. 14 km S of Comodoro Rivadavia, 24 Jan. 1968, K. A. Santarius 913 (MO-2155717, holotype; DUSS, M, isotypes). Distribution (Fig. 230): So far known only from a few stations in the prov- inces of Buenos Aires, Chubut, and Santa Cruz, Argentina. Specimens examined from cultivated plants: RGENTINA. BUENOS AIRES: Dunes about 2 km SE of Argerich in the research terrain of the Universidad del Sur, 37 km W of Bahia Blanca, Santarius 401* (CTES, DUSS, M, MO). CHUBUT: Gravelly and sandy places in Villa Balneario Rada Tilly, ca. 14 km S of Comodoro Rivadavia, Santarius 913*, 918, 923*, 924*, 925*, 930, 935, 936 (DUSS; 923 also CTES; 925 also M; 913, 923, 925 also MO). Additional specimen examined: ARGENTINA. SANTA CRUZ: Comandante Piedrabuena, O'Donell 3921 (LIL). Oenothera rivadaviae seems clearly to be a complex heterozygote, the parents of which were O. mendocinensis and O. odorata. Hybridization with these two homozygous parental species yields plants indistinguishable from them and oth- ers indistinguishable from O. rivadaviae. The sporadic distribution of this spe- cies seems to result from its independent origin following hybridization between its parents at widely separated localities. It may be separated from O. mendoci- nensis by its sparser strigillose pubescence and wider bracts, and from O. odorata by its much smaller flowers and narrower capsules. 93. Oenothera stricta Ledeb. ex Link (“striata”), Enum Pl. Hort. Berol. 1: 377. 1821. H. F. Link misspelled Ledebour’s epithet as “striata” in publishing the species.—F ics. 4, 67-72, 136-138, 220-221. Erect annual or perhaps sometimes biennial herb, rarely decumbent or nearly prostrate, forming a rosette, unbranched or with a branched main stem and side branches arising obliquely or arching upward from the rosette, 3.5-15 dm tall. Plant = strigillose, especially below, and densely to sparsely long- and short- villous as well as glandular-pubescent. Rosette leaves narrowly elliptic to ob- lanceolate, acute, gradually narrowed to the petiole or sessile and narrowly cune- ate to acute at the base, 10-25 cm long, 0.8-2.5 cm wide; cauline leaves very narrowly elliptic to lanceolate, acute, acute to rounded at the base, sessile, 18 cm long, 0.6-2.5 cm wide; bracts narrowly lanceolate to ovate, acute, ses- sile, truncate to subcordate at the base, 2-3.5 cm long, 0.7-1.5 cm wide, mostly shorter than the capsules they subtend, rarely subequal to them; leaves plane or slightly undulate at the margins, remotely or densely serrate, the teeth blunt or sharp; margins of the bracts usually reddish. Inflorescence simple or branched. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 537 JRES 186-194. Scanning electron micrographs of seeds of taxa of Oenothera se ct. 86 8 (continued ).—186. Oenothera k tu (Chile, Atacama, Jiles 2160 ).— O. arequipensis (Peru, Arequipa, Scolnik 1019).—188. O. PM i-i (Peru, Arequipa, Johnston 3556).—189. O. featherstonei (Peru, Lima, Macbride & Featherstone 270) —190. O. nocturna (Peru, Lima, ec 13).—191. O. pied subsp. pubescens (Peru, Ayacucho, Santarius 2235).—192. O. laciniata 0 pubescens (Peru, Junin, Santarius 2189). 193. O. laciniata subsp. 1 irs ited States, Missouri, Russel in 1898, MO).—194 grandis ( United States, Texas, Lindheimer 406, MO). Floral tube 2-4.5 cm long. Buds narrowly oblong to oblong or lanceolate in out- line, green or yellowish green, often flushed with red, 1.2-3 cm long, 0.3-1 cm wide; apices of the sepals erect or divergent, 1-3 mm long. Petals broadly ob- ovate to very broadly obovate, often with a red spot at the base of each one, 1.5-3.5 cm long. Anthers 5-11 mm long. Filaments 10-20 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 2.8-6 cm long. 538 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Stigma lobes 3-6 mm long. Ovary 1.3-2 cm long. Capsule 3-5 cm long, 3-4 mm thick. Seeds broadly elliptic in outline, 1.3-1.8 mm long, 0.5-0.7 mm thick. Self- pollinating complex heterozygote. Gametic chromosome number, n — 7 (ring of 14 at meiotic metaphase I). Neotype: Grown from seeds and cultivated in the Botanical Garden of Diis- seldorf, Germany, 14 Aug. 1972. Source: Chile, Prov. Concepción, airport of Concepción, end of 1959, H. Nodt (MO-2155718, holotype; CTES, DUSS, M, isotypes). Distribution (Figs. 230, 231, 235): In Chile from the province of Coquimbo to Isla Chiloe. On the east side of the Andes in the following provinces of Argen- tina: Neuquén, Río Negro and Chubut, and in the provinces of Chaco, Córdoba, San Luis, and Buenos Aires. The localities for subsp. stricta in Ecuador (Quito) and Peru (Lima) must represent stations where the plant is adventive, as is often the case on other continents. Oenothera stricta has originated as a complex heterozygote between O. odo- rata and O. ravenii. KEY TO THE SUBSPECIES l. Petals 2.5-3.5 cm long; buds 2-3 cm long, 7 = mm thick |... 23b. subsp. altissima l’. Petals 1.5-2.5(—3.5) em long; buds 1.2-1.7 cm long, 3-7 mm thick. Pubescence shaggy; bracts tow des the sssi of the stem erect; leaves remotely and bluntly serrate 23c. subsp. argentinae 2’. Pubescence not shaggy; t bracts towards the apex of the stem ae leaves mostly sharply serra 8 23a. subsp. stricta 23a. Oenothera stricta subsp. stricta.—Fics. 4, 136, 220. O. propinqua mae Nouv. Ann. Mus. Hist. Nat. 3: 343. 1835. rLEcrorvrE: Chile, Prov. Val- paraíso, rocky and sandy places along Río Quillota, 1829, Bertero 1186 (P; LE, NY W, isolectotypes). In his protologue, Spach states that he grew plants from Bertero’s seeds, but no corresponding pec ieee material been seen. brachysepala Spach, Nouv. Ann. Mus. Hist. Nat. 3: 345. 1835. rype: Chile, Prov. Val- paraiso, Quillota, Bertero 1186 (P, holotype, F cns GH, photographs). bracteata Philippi, Anales Univ. Chile 2: 394. 1862. rype: Chile, Prov. Curico, Kona pon near 0 7 2 1861, Landbeck (SGO, holotype, GH, NY and POM photographs); Lin- naea 33: 64. arguta abet Fl. Francisc.: 212. E TYPE: United States, California, Monterey Co., Monterey, July 1891, Michener (ND propinqua var. sparsiflora Philippi, il. Univ. Chile 84: 631. 1893. rype: not located. valdiviana Philippi, Anales Univ. Chile 84: 634. 1893. LEcroryrE: Chile, Prov. Valdivia, Huchuim, 27 Jan. 1887, R. A. Philippi 5 holotype, GH, NY and POM photographs). glabrescens Philippi, Anales Univ. Chile 84: 631. 1893. rEcrorvrE: Chile, Prov. Arauco, ebu, Mar. 1883, R. A. ee (SGO, s F, GH, NY and POM photographs); Munz, Amer. J. Bot. 22: 5. Onagra arguta (Greene) Small, Bull Torrey Bot. Club 23: 172. 1896. Oenothera bracteata var. glabrescens (Philippi) Reiche, Anales Univ. Chile 98: 476. 1897. O. mollissima L. var. valdiviana (Philippi) Reiche, Anales Univ. Chile 98: 477. 1897; Reiche i ile 259. 1898. ° ° ° 99 8 O. stricta Ledeb. ex Link var. 5 e Reiche, Anales Univ. Chile 98: 478. 1897; Reiche & Philippi, Fl. Chile 260. 1898 Onothera polymorpha H. Lév. race siniela n ex Link) H. Lév., Monogr. Onoth. 363. 1909; Bull. Acad. Int. Géogr. Bot. 19: 323. 1909. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 539 O. polymorpha race propinqua (Spach) H. Lév., Monogr. Onoth. 364. 1909; Bull. Acad. Int. Géogr. Bot. 19: 323. 1909. O. polymorpha race mollissima (L.) H. Lév. var. 5 (Spach) H. Lév., Monogr. Onoth. 365. 1909; Bull. Acad. Int t. Géogr. B Oenothera mollissima sullsp. propinqua (Spach) EO a ps: Mus. Univ. Zürich 58: 390. 19 O. dade: 'Pearce" Cleland, Jap. J. Genet. 43: 332. 1968. Plants 2.5-10 dm tall, the pubescence not shaggy. Rosette leaves 10-15 cm long, 0.8-1.3 cm wide; cauline leaves 6-10 cm long, 0.6-1 cm wide; bracts 2-3 cm long, 0.7-1.2 cm wide; leaves usually thicker and more sharply serrate than in the other subspecies. Internodes between the capsules 2-4 cm long. Floral tube 24.5 cm long. Buds 1.4-1.7 cm long, 3-5 mm thick; apices of the sepal 1-3 mm long. Petals 1.5-2.5(-3.5) cm long. Anthers 7-11 mm long. Filaments 10-20 mm long. Styles 3-6 cm long. Stigma lobes 3-5 mm long. Capsule 3-4 cm long, 34 mm thick. Seeds broadly elliptic in outline, 1.4-1.8 mm long, 0.5- 0.7 mm thick. Gametic chromosome number, n — 7 (ring of 14* at meiotic metaphase I). Flowering time: Northern area, October-May; southern area, October-March. Distribution (Fig. 230): In Chile from the province of Coquimbo to Isla Chiloé, and on the east side of the Andes only at Lago Nahuel Huapí in the province of Río Negro, Argentina. The localities in Ecuador (Quito) and Peru (Lima) represent escapes from cultivation, as do those on other continents. Specimens examined from cultivated plants HILE. COQUIMBO: Pisco del Elqui, Sube | in 1961* (CTES, DUSS, M, MO). Rivadavia, T 3385* (CTES, DUSS, M, MO). varPranaíso: "El Granizo” near Limache, Göpel in * (CTES, DUSS, M, MO). At the old church in eastern part of Valparaíso, Gópel in 1961 (CTES, DUSS, M, MO). CONCEPCIÓN: At Río Bío-Bío between Chiguayante and Gualquí, Mancia in 1965* en DUSS, M, MO). At Rio Bio-Bio near Concepción, Stubbe in 1960* Airport of Concepción, Nodt in 1959* (DUSS, M, MO). Bío-Bío: Fundo Paraiso! the Monte Aguila, Gópel in ii. ( CTES, DUSS, M, MO). Los 1 Qulla se; in 1962* (CTES, DUSS, M, MO). cauTin: Quepe near Temuco, at the railroad near the Río Quepe, Stubbe in 1960* (CTES, DUSS. M, MO). At the upper cur- rent of Rio Allipén, on volcanic ashes near Volcán Llaima, Göpel in 1961* (CTES, DUSS, M, O). Fundo Walker at Rio Trancura N of Volcán Villarica, Stubbe in 1960* (CTES, DUSS, M, MO). Fundo Saelzer at the S shore of Lago Villarica, Stubbe in 1960* (CTES, DUSS, M, MO). At Rio Allipén SE of Volcán Llaima, Mittak & En in 1961* (CTES, DUSS, M, MO). vALDIVIA: On dunes near Mehuín, Nodt in 1959* (CTES, DUSS, M, MO); Nodt in 1959* (DUSS). PERU. LIMA: Surco near Lima, Diers 1091* (CTES, DUSS, M, MO). Hecht 1964-8*, source unknown (CTES, DUSS, M, MO). O. stricta “Santa Barbara,” source unknown, Cleland 1967-403* (CEES, DUSS, M). Representative specimens examine HILE. COQUIMBO: Talahueri near Ovalle, Geisse in 1889 (SGO). Aconcacua: Zapallar, Behn 22800 (CONC). VALPARAISO: Quillota, Philippi in 1856 (W). Between Horcones and Ventana, Simon 290 (RSA). Limache, Looser 2008 (GH). Las Zorras, Harshberger 3391 (NY). Between Curacavi and Casablancas 600 m, Killip & Pisano (RSA, US). Mirasol near Algarrobo, Kausel 3414 (LIL). Valparaiso, so 251 (BM); Valentín 120 (S); Maximo- witsch 28 (LE); Mertens (LD, LE). Río Aconcagua near Caleta de Concón, Moore in 188 560 "rx Philippi in 1866 (SGO-059879). Vina del Mar, Philippi in 1882 (BM); Hicken 263 (SI E intero, Pühlmann in 1912 (LD). SANTIAGO: Coline a, Gay 1232 (SGO). San Antonio, ae 9 (P); 3 a 1921 (S). COLCHAGUA: Landbeck in 1860 (SGO). cuniCO: Cordillera, i Plandion, pass to Arge es Bürger in 1903 (GOET). Llico near 1 Philippi 613a (SGO). maure: Kuntze in 1892 (NY). Cauquenes, 1,000-1,650 , Ball in 1905 (K). Constitución, Reiche (SCO). LINARES: Panimávida, Philippi in 1885 O 4 m a E cen "Sg go ez M BO! 540 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 (BM); Holway 224 (US). Longavi, Schönemann in 1888 (SGO). NuBLE: St. Facelco, 520 m, Pfister 8692 (CONC). Between Curanipe and go Cox in 1962-1963 (SGO). Hacienda Los Mercedes near Chillán, Ruiz in 1995 (POM). Puente El Roble near eee Parra & Rodriguez 107 (CONC). concercién; Fundo Trinitaria near Concepción, Pfister 2166 (CONC). 5 Dombey in 1782 (P); Neger in 1893-1896 (M); Jaffuel 3995 (GH); Mertens (LE). Hualpén near Concepción, Barros 6474 E Ricardi 658 (LIL). Rio Bio-Bio, dae A lliott 86 (BM, NY). Isla Sta. Maria, Eights (US). San Pedro, Villarod 32162 (CONC). "d Ricardi 10938 (CONC). Coronel, Ochsenius in 1866 (BR, GOET ). 9 8 0 Hill 162 (K). Lota, Philippi 610 (SGO). Bio-pio: Fu + Tambillo near Nacimiento, Pfister 719 U 186 (LIL). Antuco, Poeppig in 1829 (M). ARAUCO: Arauco, Pennell 12920 (F, GH, NY, S, SGO, US). Isla Mocha, Behn 25366 (CONC). ma- LLECO: “San Lorenzo” near Angol, N. N. in 1933 (UC). Mininco, Schwabe 13338 ( CONC). Collipulli, Ricardi 7455 (CONC). Curacautin, Burkart 575 (LIL, SI). Renaico, Philippi in 1887 (860052830). caAvrIN: Rio Pedregoso near Toltén, Friedrich 3797 (CONC). Rio Zuapa, Middleton in 1905 (BM). Bajo Imperial, Middlet ton in 1906 (BM, G). Gral. Lopez near Temuco, Sandeman 346 (BM). Temuco, Claude-Joseph 1843 (US). vaArpivia: Philippi in 1888 (K); Calvert in 1914 (BM); Buchtien in 1900 (US). San Juan, Philippi in 1865 (SGO-052881). Niebla near Valdivia, Buchtien in 1899 (US) (as O. mollissima var. sabulosa Buchtien). Ranco, Philippi in 1887 (SGO-052860). Futronhue, Philippi 613d (SGO). Cerro Llifén, Marticorena 63 (CONC); Boelcke 254 (SI). Bellaviste in the valley of Rio Trumas, Lechler 416 (GOET, K, P). cuiLoÉ: Chacao, Bartulin 12462 (CONC). Isla Chiloé, Ruiz POM); Nr. 126 (SGO-052883). Castro, Philippi in 1880 (SGO-052862). Río Palena, Delfin in 1887 (SGO). 2GENTINA. RIO NEGRO: Lago Martin Steffen, Boelcke 6236 (BAA). Nahuel Huapi, Arroyo Los Cornelios in the valley of Rio Limay, Boelcke & Hunziker 3628 (BAA, MO). BUENOS AIRES: Bahia Blanca, near the port, Reineck in 1899 (L) (adventive ). Ecuapor. Parque de Ibarra , 2,225 m, 1949, Solis 13390 (F). Peru. Rimac valley near Lima, 700 m, 1954, Rauh-Hirsch 115 (RSA). Specimens from outside of S outh America: UNITED STATES. CALIFORNIA: Monterey Co., Heller in 1903 (MO). Mexico. College garden at Morelia near Michoacán, Arséne in 1908 (Z). Hawai. Maui, Olinda, Degener in 1927 (MO). Hawaii, Haleakala crater near Holua caves, 1927, Degener 2254 (K). keo ca. 2,450 m, 1930, John 10341 (K); 1940, Mebold 26664 (M). Kilauea, 1935, Mahali 20960 (M). ScorLAND. Skin works near Selkirk, ion Hayward 1060, 1913, 1065 (E); 1966, Web- ster 10959 (E). Wares. Aberdovy, Melville in 1919 (BM), 1919, Melville 48 (E); 1928, Britton 3390 (K); 1948, Taylor 1342 (K). Glamorgan, Cumming in 1922 (BM, K). somerset: Duthie in 1869 (BM). Burnham, Melville in 1873 (E); Thompson in 1898 (BM, E, K); Fogitt in 1932 (BM); Alston in 1949 (BM). Warwick, Bromwick in 1870 (BM). Berrow, Davis in — — ~ — BM). ND. BEDFORD: Patton, 1950, mie a = 1966, Webster 10942 (E). CHESHIRE: Bickle 5 peer d in 1902 (BM). CORNWALL: Gw Davey in 1903 (E). Par, Medlin in 1920 (K); Mebold in 1924 (M). c Druce i in 1926 (BAS). pEvoN: Laira near icq outh, e in 1863 (BM). Dawlish Warren, Redgrove in 1937 (BM); Proctor in 1952 (LISE). DORSET: rcham, Fawett in 1884 (BM); Linton in 1893 (BM). HAMPSHIRE: l 4 Alston in "n (BM). Blackmoor, 1966, Webster 10900 (E); 1968, Lousley 3207, 3262 (BM KENT: Catford, Lowne in 1910 (K). Sandwich Bay, Meinertshagen in 1932 (BM); Town- send in 1948 (K); 1961, Butler 268 (BM). Castle near Sandwich, Reid in 1975 (BM). Rich- borough, Mill in 1860 (K). MIDDLESEX: Twickenham, Clifton Road, in 1867 (BM). Wands- worth in London, Forbes in 1837 (BM). NORFOLK: Jarmouth, Linton in 1879 (BM, K); Bichham in 1901 (K). surrey: Croydon, Bennett in 1873 (E). Bisley Railway, 1958, Burkill 1639 (E). IsLe or wicur: Fawett in 1879 (BM); Jackson in 1894 (K); 1933, Koster 775 (K); Townsend in 1951 1 CHANNEL ISLANDS: Guernsey, L’Ancresse in 1884 (BM); Balfour- Brown in 1950 (BM). Jersey, Watson in 1864 (BM); Gray in 1894 (BM); 1920, MacAlister Hall — (E); 1954, Duncan 775 es RELAND: WEXFORD: Rosslare, Druce in 1926 (BM). 5 VIENNE: Chatellerault, Chabaipeau in 1860 (K, P). LOIRE-ATLANTIQUE: Bour- neuf, Gad. a 1896 (BM). VENDÉE: Challans, Portineaux in 1897 (BM). L'Aiguillon, Fou- aud i 9 (LY); 1911, Hibon 1461-3 (P). GIRONDE: Bégles, Neyraut in 1892 (MPU). Pauillac, 1 Jallu 461 -4 ( (MPU). LANDEs: Capbreton, Foucaud. in 1880 (LY); 1913, Hibon 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 541 1461-3 (P). Parentis-en-Bors, 1962, Retz 49626 (LISE). BASSES PYRENEES: St. Jean de Luz, Barbey in 1883 (Z). Bayonne, 1883, Blanchet 546 (LISU, LY, MPU, P). Jetée de Boucaud, Foucaud in 1883 (LY); Fourés in 1909 (MPU). AVEYRON: Villefranche, Bras in 1882 (P). HERAULT: Castelneau near Montpellier, Barrandon in 1858, in 1862 (BAS). Séte, Neyraut in 1887 (MPU); Cabans in 1933 (MPU). var: La Garde- Freinet, Défends du Refréne, 1900, Bertrand 4725 (HBG, LY, MPO, P), in 1901 any. in 1903 (LY). Grinaud, Hibon in 1920 ). Île d'Aurigny, Corbière in 1888 (M). "ag Normandie, St. Sauveur- de-Pierrepont, LY). Corbiére in Ses (LY). FINISTÈRE: Bretagne, coff, Miciol in 1890 ( CHARENTE MARITIME: endre, Foucaud in 1876, 1897 a 1898 (LY). For hybrids with Oenothera longiflora 1 9 pei. see p. 514. TALY. Viareggio, Ball in 1866 (K); Caldesi in 1880 (LD); Lévier in 1882 (P); Gibelli in 1886 (GOET, LY, MPU); Mori in 1886 (LD); Knetsch in 1903 (Z); 1908, Fiori 1316 ( BM, E, K, LY, Z). PORTUGAL. Figueira, Mariz in 1882 (COI); 1885, Goltz de Carvalho 825 (COI, LISE, LISU); Costa in 1933 (COI); Silva in 1940 (LISE); Matos in 1948 (C); 1966, Reis Moura 706 (COI). Ovar, 1951, Silva et al. 4563 . 5 1966, Merxmiiller & Grau 21497 (M). Caldas da Rainha, Murray in 1889 (BM). Moita, da Cunha in 1882 (LISU); Jorge & Mendez in 1917 (LISU); 1942, da Silva 140 (LISE); 1954, Rainha 2762 (LISE); Fernandes et al. in 1961 (COI, LD). Sintra 1944, Rainha 46, 1950, 1983 (LISE). Martiganca, 1956, da Silva et al. 5727 (LISE). Colares, da Cunha in 1943 (LISU). Cuitra, dos Santos in 1909 (LISU ). Rocha, Choffat & Daveaux (MPU). Mizarella near Guarda, Ferreira in 1885 (COI). Villar Formoso, Mariz in 1900 (COI). Marinhais, 1946, Garcia & Souza 918 (COI). DEIRA. Cerro de S. Roque, Mandon in 1865-1866 (G, K). IN. Gigón, Bourgeau in 1864 (C). Lá Coruña, Guardi in 1888 (BM). San Sebastian near Guipuzcoa, 1895, Gandoger 186 (C, COI, K). Pr rov. Gaditana, near Linea de Concep- ción, 1895, Porto & Riga 639 (B). Gibraltar, Linea sand hills, Wolley- Dod in 1912 (BM). Almoreima near Algeciras, 1924, Ellman & Hubbard 631 (K). Puerto de Santa María near Cadiz, 1929, Ceballos 2184 (BM, Z); 1968, Merxmiiller & Lippert 23327 (M). Banks of Rio deus i in Andalusia, 1955, Brinton Lee 87 (BM). GERMANY. Port of L Ludwigshafen, Zimmermann in 1910 (BAS). Kettwig, Bonte in 1912 (BA s A near Basel, Herb. Fischer (M). European part of USSR, spontaneous in 1964 in a potato field belonging to the 2 Garden of the University of Moscow, Skortsov 10188 ( 1 MHA). Not to be regarded as a regular member of the adventive Hoe of European Russ Japan. Cultivated, Yokohama, in 1862 (BM). Kir, 1875, Rein 20 (COET). Kyoto, Rein (BM, HBG, M); Hikko in 1877 (HBC). Tokio, rns Drake in 1881 (P). Harima near Hondo, Arimoto in 1903 (MO). Kyoto, 1907, Dunn 8706 (K). Aboshi, 1912, Schwarz 0 (Z). Honshu, Ashiya, Fox in 1912 (BM). Kiushu, Hakozaki near Fukada, 1928, Ichi- kawa 62 (BM, P). Honshu, Imazu river, 25.6 mi W of Hiroshima, Charette in 1953 (MO). Niigata, Drake 52 (P). Yokoka, Drake (P). AKISTAN. North-West Himalaya, Murree, Saunders in 1915 (K); 1918, Stewart 4039 (RAW INDIA. Nilghiri, Ba Mes 408 (P); 1850, Hohenacker 1146 (K); Gamble in 1886 (K). Ootacamund near E . 2,300 m, 1882, Brandis 551 (HBG). Punjab, near Bana- sar, 1885, Drummond 24435 (K). Punjab, 1922, oe 24434 (K). Himalaya, at the railway between Suni hill and Tulogk, Rich in 1916 (K ashmir, vicinity of Dalhourie, ca. 2,350 m, 1917, Stewart 2140 (K). Almora, 1,560 m, 826 a, 2021, 2128 (K). Between Kodai Channel and Pulneys, Bourne in 1898, 1899 (K); Foreau in 1960 (K). CrYLon. Roadside near Ambawella, 1932, Simpson 2 BM). Java. 1915, N.N. 107, 120 (K). Tjibodas, Mt. Pangrango, rare, 3,000 m, 1948, Djambari 317 (L); 1950, van Ooststroom 13339 (L). USTRALIA. WESTERN AUSTRALIA: Lower Swan River near Bayswater, Morrison in 1907 M). Bayswater, Howard 231 (K). Katanning, Dowell in 1954 (PERTH). Woodman’s point, 1961, Aplin 1071 (PERTH). Claremont, Steward in 1961 (NSW-135207). Nicholson Road near Connluten. 1961, George d Marchant 3155 (PERTH). 19 mi SE of Nyabing, 1962, Newbey 400 (PERTH). Between Borden and Abbany, 1965, Anway 573 (PERTH). Pian Spercer & Fievez in 1971 (PERTH). SOUTH AUSTRALIA: Murrumbidgee, Nolan in 1841 (MEL). Port Elliot, Hussey in 1893 (MEL). Mt. Lofty near Adelaide, in 1897 (MEL). Murray Bridge, Maiden in 1907 (NSW-135206). Adelaide, Kaspiew in 1951 (BREM). Wiltunga on Northern York Peninsula, 1966, Copley 114 (K). Mt. Lofty Range, Balhannah 25 km SE of Adelaide, 1966, Eichler 18902 (K). Mt. Gambier, 1966, Wilson 646 (C, MEL). 542 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 W shore of Lake Albert, 14 mi W of Meningie, pé Willis 622 (MEL). vicronta: Upper Wimmera River near Stawell, Matthews in 1893 (MEL). Wangaratta, Weir in eds ( MEL). Wimmera, Walter ( MEL-58120). 4 mi NW of Shepparton, between Angustown and Wahoo Ackland in 1963 (UPS). 5 mi W of Ararat, 1964, Muir 3390 (MEL). Red Chi Henshall in 1964 (MEL). P 5 11.7 mi N of Hopetoun, 1968, Belcher 1624 (MEL). SOUTH WALES: Balnarand, 1878, Lucas 75 (MEL). Filba, Reader in 1880 7 5 Riverside near Coonabarralaan, L. in 1883 (BM). Hume River, Scott in 1883 (M m Turon River, 1885, Lauterer 40 (MEL). Wagga, Fletcher 2 1888 (NSW-135215). Kogarah, Campfield in 1893 (NSW-135238). Springfield, 25 mi SW of Cobar, Andrae in 1895 (MEL-58123). Barbes Creek near Tallong, Rumsey in 1898 Tous 135240). Jenolan Caves, Blakely in 1900 (NSW-135231). Orange, Boorman in 1906 (NSW-135236). Bega, Ziethen in 1906 ( NSW- 135228). Richmond, Carne in 1906 (NSW-135242). Blackheath, Maiden in 1908 (NSW- 135233). Emu plains, Hamilton in 1912 (NSW-135241). Wallendbeen, Beakwell in 1913 (NSW-135219). Bathurst Farm, Noble in 1914 (NSW-135235); Brett in 1914 (NSW-135234). Hill TOP Southern Line, Cheed in 1915 (NSW-135237). Tinga, Boorman in 1917 (NSW- 135222). Blue Mountain, N.N. in 1919 (MEL-58109). Rylstone, 1920, Morton 5865-20 NSW). Tumut, poda in 1921 (NSW. 35214. Cocketgedong near Jerilderie, Sibb in 1921 (NSW-135224). Wondabyne to Woy Woy, Blakely in 1922 ( NSW-135243). Junee Distr., Nugent in 1993 (NSW-135216). Harden, Rodway in 1924 (NSW-135218). Wallangra, Rod- way in 1929 (NSW-135223). Cullerin, 25 mi W of Goulburn, Simpson in 1935 (NSW- 135229). Armidale University Grounds, Davis in er ( NSW- 1523 30). Morivale to Welling- ton, Shelley in 1946 (NSW-135212). Narrabeen Lake, Johnson in 1946 (NSW-135239). Carlaminda, Castin in 1948 ( NSW-135227). Near Wellington, Dunk in 1950 (MEL-58103). Narrhari, Moore in 1952 (NSW-134226). Albury distr., Glenfield in 1953 (NSW-135213). Near Kootingal, 8 mi ENE of Tamworth, 1954, Kelso & Goode 64 (NSW-135220). Menindee, Constable in 1955 (K, MO, NSW-38446). Near Wyong, 1958, Salasoo 1633 (NSW). Cowro Research Station, Hill in 1960 (NSW-135210). Canberra, 1962, Mukee 9666 (NSW). Coota- mundra, diae in 1963 (NSW-135217). Narrandera, Leeton in 1963 (NSW-135225). Gun- nedah Distr., Beeson in 1964 ( NSW-135221 ). Michelago to Williamsdale, Salasoo in 1969 (NSW). Bylong. 33 mi NE of Mudgee, 800 m, 1970, Raven et al. 25868 (MO, NSW). pes River, 39 mi SSW of Jindabyne, 1970. Pickard € Coveny 2758 (MEL, NSW). QUEENSL Caves in 1874 (MEL). Silverwood, 1922, White 1735 (NSW-135209). Mt. Playfair sete near Leichhardt, 1964, Adams 1352 (K). Maryland, Hickey in 1884 (MEL). Boxbill, Reader in 1884 (MEL). Lilliput, in 1913 (MEL). Near Klunzy, Mueller (MEL-58115). TASMANIA: Hobart, Lucas in 1913 (NSW-135208). NEW ZEALAND. Roturoa, Chase et al. in 1909 (BM, LIL, K, MO). Swamp near head of Rangaunu, harbour of Puheke Hill, Mason & Moar in 1949 (Al). Sulphur Springs Bay near Rotoity, Hurvey in 1949 (Al). Owairaka Park Domain, Wood in 1950 (AI). Coal Creek, 90 Mile Beach, Cooper in 1966 (AI). Murimotu at Nort h Cape, Adams in 1968 (AI). New Zealand, Hooker in 1870 (K). Paumure, TFC (AI). LipyA. Distr. Tripolis, Sidi Mesri, 50 m, 1960, Keith 827 (K). Ecvrr. Near a small village at the Channel in 1872 (BM). RuopEsia. Salisbury, on railway, ca. 1,650 m, 1919, Eyles 1533 (BM, SRGH). Dist Mtoko, ca. 1,000 m, 1956, Davies 1926 (K, SRCH). Melsetter village, 1968, Goldsmith 126. 68 (M, SRGH); 1960, Phipps 2844 (SRGH). MOZAMBIQUE. Inhaca Island, 23 mi E of Lourenço Marques, 1959, Mogg 31640 (K). SOUTH AFRICA. CAPE: Loerie, 1894, Penther 2150 (M). Distr. Aliwal North, Kraai River, ca. 1,450 m, 1933, Gerstner 198 (PRE). Distr. East London, 1960, Comins 2038 (PRE). Distr. Alexandria, Reed Valley, ca. 320 m, 1953, Archibald 5886 (PRE). Distr. Humansdorp, 1921, Fourcade 1762 Somerset, Bowhen (K). Stellenbosch, 1948, Parker 4391 (K). Rondebosch, 1934, Adanson 2232 (BM). TRANSVAAL: Woodbush, 1909, Jenkins 7201 (PRE). Bokfontein, 1909, Jenkins 7542 (PRE). Witzies Hoek, 1917, Junod 14522 (PRE). Grahams- town, 1918, van Dam 18837 (PRE). Knysna, 1921, Breyer 28334 (PRE). Along railway near Benoni, 1934, Bradfield 189 (PRE). Pretoria, 1936, Reptar 650 (PRE). Bethlehem, Potgieter 21840 (PRE), ORANGE FREE STATE: Clarence, 1917, Van Hoepen 18172 (PRE). Gansfontein near Ficksburg, ca. 1,750 m, 1934, Galpin 13877 (K, PRE). Doornhoek near Bloemhof, Botha in 1936 (PRE-29489). Bloemfontein, 1951, Gemmell 6142 (K, PRE); Hane- kom 811 (K, SRGH); 1951, Potts 6142 (K). Natal: 1902, N.N. 41 (BM). tesoro: Distr. Tebetebeng, 1957, Jacot- Guilumod 2928 (PRE). Maseru, ca. 1,650 m. 1970, Williamson 19 (K). Leribe, Dieteren 169 (P). Province unknown: Wit pootje Kloof, 1948, Moss 4648 (BM). 7. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 543 Hybrids between O. stricta subsp. stricta and O. indecora subsp. bonariensis occur in T i s and Portugal, and are cited under O. indecora (p. 522). den ens: Botanical Garden of Dresden, Gern wr Bauer in 5 (CORD). Copenhagen, Herb. 1820, Schum. 18 (C). Copenhagen, Herb. 1822, Schum. 26 (C; as O. fraseri). Copenhagen, Herb. 1822, Schum. 25 (C; as O. salicifolia). Botanical 1 5 of Frankfurt, Germany, seeds from Berlin, Becker in 1825 (FR; as O. striata). Paris, in 1842 (BR). Leningrad. in 1848 (LE). Vienna, Austria, in 1849 ( W; as O. chilensis). Oenothera stricta subsp. stricta reported in literature br outside of South America: ENGLAND. Trimen & Dyer (1869: 111, as O. odorata); Babbington (1881: 136, as O. odorata); Murray (1896: 154, as O. odorata); Hanbury & Marshall (1899: 158, as O. odorata ); Linton (1900: 105, as O. odora ta); Marquand (1901: 363, as O. odorata); Lester-Garland (1903: 113, as O. odorata); Davey (1909: 203, as O. odorata); Trow (1911: vol. 1: 71, as O. odorata); Hooker Xd 158, as O. odorata ); 17 555 (1953); Butcher (1961: vol. 1: 798); Clapham et al. (1962: 480); Perring & Walters FRANCE. Gandoger (1886: 49); Corbiére Ld. 239); Burnat 1 vol. 198); Rouy & Camus (1901: 201); Coste (1903: vol 2: 81); Thellung (1912, as O. 1 subsp. odorata); Bonnier Sim vol. 4: 29); Chassagne (1957: vol. 2: 160); Issler et al. (1965: 357); Raven (1968) GERMANY. Raven (1968). e (1968). PortucaL. Coutinho (1913: 426; 1939: 508); Sampaio (1946: 408); Raven (1968). SPAIN. Willkomm & Lange (1880: 181); Willkomm (1893: 219); ner (1886: vol. 2: 389); Menezes (1914, as O. odorata); Wolley-Dod (1914: 42; 1949: IrALY. Arcangeli (1882: 238); Gandoger (1886); Fiori & Paoletti L3 vol. 2: 134); Saccardo (1909: 176); Fiori (1925: vol. 2: 14). Zangheri (1 ae vol. 1: U.S.S.R. Shteinberg (1949: 630; 1974, Engl. transl.: 472, s O. ds aven (1968). Monocco. Jahandiez & Maire (1931; 516 ALGERIA. Quézel & Santa (1963: wes SEM Bizzari & Raven (1972: 4 OUTH AFRICA. Adamson & Salter e 606); Guillarmod (1971: 215, as O. longi- lios). Ross (1972; 262). INpiA. Graham (1839: 75, a T „ Hooker (1879: vol. 2: 582, as O. odorata); Fyson (1915: vol. 1: 161, vol. 2: odorata); Trimen (1931: 131, as O. odorata). CEvLON. Trime n (1894: 23 E as T eis JAPAN. Makino (1949 290, tab. 868). CHINA. Iconographia (1972, as O. odorata). AUSTRALIA. Black (1909: 63; 1926: vol. 3: ea + O. odorata; 1952: vol. 3: 638); Beadle et al. (1962: 173, as O. odorata): Eichler (1965: 243). New ZEALAND. Kirk (1899: 180, as O. 0 Cheeseman (1925: 1073). Unirep States. Munz (1965). The stations for this subspecies at Lago Nahuel Huapi and the Rio Limay in Argentina might represent introductions, like those in Ecuador and Peru, espe- cially since there is a constant flow of tourists from Chile to the national parks in Argentina. This subspecies is morphologically similar to O. ravenii subsp. chilensis, itself an entity which includes genes from the other parent of O. stricta, O. odorata. Oenothera stricta can be distinguished from O. ravenii subsp. chilensis, how- ever, by its longer bracts and less erect habit. In fact, considering the range of variation in the species as a whole, the epithet “stricta” is a little unfortunate, A. K. Skvortsov has kindly informed us that the specimen (Skvortsov 10188, DS, MHA) on which Raven (1968) based his record from European Russia was from adventive plants which appeared only in 1964. The species is naturalized in the vicinity of Vladivostok. 544 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 195 196 197 198 199 200 201 202 203 204 205 206 207 208 209 Ficures 195-209. Schematic outlines of buds id taxa of Oenothera sect. Oenothera sub- sect. Munzia ae he santarii (Argentina, Mendoza, Santarius 1495).—196. O. longituba (Argentina, Tucumán, Santarius 1736).—197. O. 3 (Argentina, Jujuy, Fabris 5787). —198. O. 5 (Argentina, Tucumán, Santarius 1781) —199. O. scabra (Bolivia, Cochabamba, Santarius 2003). —200. O. scabra (Peru, Ayacucho, Santarius 2251).— 20 : ri u x 205. O. villaricae (Chile, Valdivia, Göpel in 1961).—206. O. cordobensis (Ar nimus. LN Gópel in 1961 ).—2 O. acuticarpa (Argentina, Tucumán, Göpel in 1961 ( Argentina, Dr Santarius 1742).—209. O. verrucosa (Peru, Arequipa, 1 2073). since there are decumbent or nearly prostrate dune and strand forms, such as Nodt in 1959 Especially in the northern part of the range, in the provinces of Coquimbo and Valparaiso, plants with a relatively long floral tube (44.5 cm) are frequent (e. g., Gdpel in 1961, Valparaíso). They correspond more or less to the type of O. brachysepala Spach. The most likely explanation for the origin of these plants seems to be introgression from O. affinis, either directly or via O. picensis, which contains one complex from O. affinis and like it grows with O. stricta in northern Chile. 23b. Oenothera stricta subsp. altissima Dietrich, subsp. nov.—Fics. 67-69, 137, Plantae 5-15 dm altae; pubes non hirta. ey Mee 10-20 cm longa, 1.2-1.7 cm lata; = apnea 7-15 cm longa, 1-17 cm lata; bractea 2-3.5 cm longa, 0.5-0.7 cm lata; folia remote obtuseque serrulata. Internodia inter ee 4-5 cm longa. Tubus floralis 2.5-3.5 cm ea Gemmae 2-3 cm longae, 7-10 mm crassae; apices sepalorum 1-3 mm longi. Petala cr 1.8 mm longa, 0.6-0.7 mm crassa. ances gameticus chromosomaticus, n = 7; planta chro- mosomatice S ed complex 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 545 Plants 5-15 dm tall, the pubescence not shaggy. Rosette leaves 10-20 cm long, 1.2-1.7 cm wide; cauline leaves 7-15 cm long, 1-1.7 cm wide; bracts 2—3.5 cm long, 0.5-0.7 cm wide; leaves remotely serrate, with dull teeth. Internodes between the capsules 4-5 cm long. Floral tube 2.5-3.5 cm long. Buds 2-3 cm long, 7-10 mm thick; apices of the sepals 1-3 mm long. Petals 2.5-3.5 cm long. Anthers 6-7 mm long. Filaments 13-18 mm long. Style 3.5-5 cm long. Stigma lobes 3.5-6 mm long. Capsule 3.5-5 cm long, 3-4 mm thick. Seeds broadly ellip- tic in outline, 1.5-1.8 mm long, 0.6-0.7 mm thick. Gametic chromosome number, n = 7 (ring of 14* at meiotic metaphase I). Flowering time: November-March. Type: Grown from seeds and cultivated in the Botanical Garden of Düssel- dorf, Germany, 11 Aug. 1972. Source: Argentina, Prov. Río Negro, NW slopes at Ruta 258 ca. 3 km N of Río Villegas (67 km S of San Carlos de Bariloche), on volcanic ashes and marl, 700 m, 19 Jan. 1968, K. A. Santarius 798 (MO-2155212, holotype; CTES, DUSS, M, isotypes). Distribution (Fig. 231): Along the eastern side of the Andes at low eleva- tions in the provinces of Neuquén, Río Negro, and Chubut, Argentina. Specimens examined from cultivated plants: ARGENTINA. PROV. RÍO NEGRO: NW slopes S Ruta 258 ca. 3 km N of Río Villegas, 700 m, Santarius 798*, 799* (DUSS, MO; 798 also M). N slopes at Ruta 258, 1.5 km S of Río Ville gas, on volcanic ashes and marl, 550 m, Santarius 800* (DUSS, M, MO). Additional specimens examined: ARGENTINA. NEUQUÉN: Cascada Maipü, De Barba 1839 (LIL). Lago Lolog, De Barba 1794 (LIL). Aluminé, Soriano 1272 (BAB). Lago Epulafquén, Dawson d» Schwabe 2456 (BAB). San Martín de los Andes, De Barba 1694 (LIL); O'Donell 2373 (LIL, NY), 2415 (LIL); Eskuche 01329 (CTES). Quila Quina, Cabrera 20521 (LP, P); Achajonsky 3399 (BAB). Lago Nonthué, Hua Hum, Valla et al. (MO). Chos Malal, ca. 650 m, Comber 169 (K). nio NEGRO: S.C. de Bariloche, De Barba 322 (LIL). Cerro Granito near Bariloche, Meyer 8244 (LIL). At Ruta 258 near Río Foyel, Dawson 3290 (BAB, LP). Villegas, Lour- teig & Buchinger 201 (P); Moreau in 1941 (RSA). El Bolsón, Meyer 7884 (LIL). cHupur: Epuyén near Cushamén, Muniez 5504 (BAB). Rio Corcovado, Illin 147 (UC). Considering the relatively large size of its flowers, O. stricta subsp. altissima may be at an earlier stage in the evolution of complex heterozygosity than is subsp. stricta. Since the influence of O. odorata is predominant in the phenotype of O. stricta subsp. altissima, one could assume that this entity arose from the combination of a chromosomally homozygous strain of O. odorata, common in the right area, with a genome of O. ravenii introgressed with O. odorata, the latter probably by way of the Chilean subspecies of O. stricta. The direct com- bination of O. odorata with the homozygous, large-flowered O. ravenii seems very unlikely, especially since the two are not known to grow together anywhere. 23c. Oenothera stricta subsp. argentinae Dietrich, subsp. nov.—Fics. 70-72, Plantae 5-13 dm altae; pubes hirta. Folia rosulae 10-25 cm longa, 1.5-2.5 cm lata; folia caulina 7-18 cm longa, 0.8-2.5 cm lata; bractea 2.5-3.5 cm longa, 0.7-1.5 cm lata, ad apicem caulis aliquantum imbricata; folia remote obtuseque serrata. Internodia inter capsulae 1.5-3 cm longa. Tubus floralis 2-3.5 cm longus. Gemmae 1.2-1.5 cm longae, 5-7 mm crassae; api- ces sepalorum 1.5-2 mm longi. Petala 1.5-2 cm longa. Capsula 3-4 cm longa, 3-4 mm lata. 546 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 210 21 212 213 214 215 216 27 28 219 220 221 222 223 224 FicurEs 210-224. Schematic outlines of buds of taxa of Oenothera sect. Oenothera sub- sect. Munzia.—210. O. mendocinensis (Argentina, Buenos Aires, Santarius 411).—211. O. odorata (Argentina, Neuquén, Santarius 694).—212. O. ravenii subsp. ravenii (Brazil, Rio Grande do Sul, Hackbart in 1966).—213. O. longiflora subsp. grandiflora (Argentina, Corri- entes, Krapovickas & Cristóbal 11293).—214. O. longiflora subsp. longiflora (U bia Colo- nia, Santarius 82).—215. O. indecora subsp. indecora (Uruguay, Florida, Santarius 212) — —217. O O. indecora subsp. indecora 11 Florida, Santarius 20 ; Don subsp. bonariensis (Botanical Garden Erlangen).—218. O. affinis (Argentina, E oe 1851). O. mollissima (Uruguay, ee Santarius 32).—220. O. stricta subsp. stricta (Chile, Concepción, Stubbe in 1960).—22]. O. stricta subsp. ‘altissima 1 1 Rio Negro, Santarius 800). —222. O. picensis subsp. cordobensis (Argentina, Córdoba, Cöpel in 1961). 223. O. parodiana subsp. parodiana (Uruguay, Florida, Santarius 207 ). —224, O. nocturna (Peru, Lima, Santarius 2333). Semina aue late elliptica, 1.3-1.5 mm longa, 0.6-0.7 mm crassa. n gameticus chro- mosomaticus, n = 7; planta chromosomatice heterozygotica complex Plants 5-13 dm tall, the pubescence shaggy. Rosette leaves 10-25 cm long, cm wide; cauline leaves 7-18 cm long, 0.8-2.5 cm wide; bracts 2.5-3.5 cm long, 0.7-1.5 cm wide, overlapping to a considerable extent towards the apex of the stem; leaves remotely serrate, with blunt teeth. Internodes between the capsules 1.5-3 cm long. Floral tube 2-2.5 cm long. Buds 1.2-1.5 cm long, 5-7 mm thick; apices of the sepals 1.5-2 mm long. Petals 1.5-2 cm long. Anthers 5-7 mm long. Filaments 10-12 mm long. Style 2.8-3.7 cm long. Stigma lobes 3-6 mm long. Capsule 3-4 cm long, 3-4 mm thick. Seeds broadly elliptic in outline, 1.3-1.5 mm long, 0.6-0.7 mm thick. Gametic chromosome number, n = 7 (ring of 14* at meiotic metaphase I). Flowering time: November-April. 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 547 Type: Grown from seeds and cultivated in the Botanical Garden of Düssel- dorf, Germany, 14 Aug. 1972. Source: Argentina, Prov. Buenos Aires, E end of the Sierra del Volcán NW of Puerta El Abra, at km 45 of Ruta 226 between Mar de la Plata and Balcarce, partly grazed terraces between rocks, 150-300 m, 7 Jan. 1968, K. A. Santarius 346 (MO-2155215, holotype; CTES, DUSS, M, isotypes). Distribution (Fig. 235): Known only from a few localities in the provinces of Chaco, Cérdoba, San Luis, and Buenos Aires, Argentina. Specimens examined from cultivated plants ARGENTINA. BUENOS AIRES: E end of the Siena del Volcan, at km 45 of Ruta 226 between Mar de la Plata and Balcarce, 150-300 m, Santarius 346*, 347, 348, 350-354, 355*, 356-367, 368* (DUSS; 355, 368 also CTES; 346, 355 also M; 346, 352, 355, 358, 368 also MO). Additional specimens examinec ARGENTINA, CHACO: Colonia Bouts Schulz 792 (POM). Las Palmas, Jórgensen 2487 (LIL). corpospa; La Isla near Santa María, de la Sota 708 (LIL). La Falda near Punilla, Stuckert 4323 (CORD). Casa Grande near Punilla, de la Sota 3411 (LIL). Cruz Grande near Punilla, de la Sota 3634 (LIL). san Luis: Paucanta, Castellanos 25-864 (POM). BUENOS AIRES: S. Vigilancia near Balcarce, Jurado 140 (LIL). This subspecies can be distinguished from the other two by its shaggy pubes- cence, a characteristic that is often found in O. ravenii. The inflorescence of this subspecies is more crowded than in the others, and the internodes between the capsules shorter. It is more heavy-set than in the other subspecies, with the thicker stems and broader leaves especially prominent. In all of these respects, O. stricta subsp. argentinae approaches O. ravenii, which predominates in its genetic makeup in the same way that O. odorata does in that of O. stricta subsp. altissima, a relationship that can easily be understood on geographical grounds. 24. Oenothera bahia-blancae Dietrich, sp. nov—Fics. 73-75, 139. erba annua erecta, rosulata, simplex n caulis principalis ramosus et ramis oblique e dade. pud ase di. 5-8 dm alta. Plantae dense vel sparse strigulosae, pilis brevibus longibus- que sparse praeditae, et sparse fandili oso- pulse Folia 5 8 anguste oblanceolata, acuta, lamina in petiolum gradatim decrescens, 10-15 cm are. 0.5-1.2 cm lata; folia caulina emas m ey en lanceolata, acuta, sessilia, basi anguste cuneata vel acuta, 5- 10 cm lon 0.6— n lata; bractea anguste 1 s a vel Proclus, Dor sessilia, basi rot es TA Sid truncata oe 3 cm longa, 0.7-1 cm lata; folia plana vel ad margines undulata, irregulariter obtuseque . Inflorescentia simplex vel ramosa. Tubus floralis 1.2—2.5 cm longus. Gemmae bino oblon ng = vel lanceolatae, virides vel flavovirescentes, saepe junctura .6-1.2 c —4 mkeo punctata; ere RUE erecti, 1-2 mm longi. Petala obovata vel latissime obovata, is, sti 1 1 5 cm longum. Capsula (2) 34.5 cm longa, 3-4 mm crassa. Semina ambito late elliptica, 1.2- 5 mm longa, 0.6-0.8 mm crassa. Numerus gameticus chromosomicus, n = 7; planta chromo- somatice heterozygotica complexa. Erect annual herb, forming a rosette, with a simple or branched main stem and the side branches arising obliquely from the rosette, 5-8 dm tall. Plants densely to sparsely strigillose, sparsely long- and short-villous, and sparsely glandular-pubescent. Rosette leaves narrowly oblanceolate, acute, gradually nar- rowed to the petiole, 10-15 cm long, 0.5-1.2 cm wide; cauline leaves narrowly elliptic to narrowly lanceolate, acute, sessile, narrowly cuneate to acute at the base, 5-10 cm long, 0.6-1.5 cm wide; bracts narrowly lanceolate to lanceo- late, acute, sessile, rounded to truncate at the base, 2-3 cm long, 0.7-1 cm wide; 548 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 $00 MILES — ° 200 400 800 800 KILOMETERS ` 60 so 40 30 20 ——— S n 9° 60 225. Ranges of Oenothera peruana (dots), O. lasiocarpa (circles), O. santarii (filled 5 and O. ravenii subsp. ravenii (hollow triangles). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 549 leaves plane or undulate at the margins, irregularly serrate, the teeth blunt. Inflorescence branched or unbranched. Floral tube 1.2-2.5 cm long. Buds ob- long to lanceolate in outline, green to yellowish green, often red striped at the junction of the sepals with the floral tube, 0.6-1.2 cm long, 3-4 mm thick. Sepals often flecked with dark red; apices of the sepals erect, 1-2 mm long. Petals obovate to very broadly obovate, 0.7-1.5 cm long. Anthers 4-6 mm long. Fila- ments 5-10 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 1.8-3.2 cm long. Stigma lobes 3-4 mm long. Ovary 1.3-1.5 cm long. Capsule (2-)3-4.5 cm long, 3-4 mm thick. Seeds broadly elliptic in outline, 1.2-1.5 mm long, 0.6-0.8 cm wide. Self-pollinating complex heterozy- gote. Gametic chromosome number, n — 7 (ring of 14* at meiotic metaphase I). Flowering time: November-April. Type: Grown from seeds and cultivated in the Botanical Garden of Düssel- dorf, Germany, 14 Aug. 1972. Source: Argentina, Prov. Buenos Aires, dunes ca. 2 km SE of Argerich in the research terrain of the Universidad del Sur, 37 km W of Bahía Blanca, 9 Jan. 1968, K. A. Santarius 455 (MO-2155721, holotype; CTES, DUSS, M, isotypes). Distribution (Fig. 236): Known only from the provinces of Buenos Aires, La Pampa, Neuquén, and Chubut, Argentina. Specimens examined from cultivated plants: ARGENTINA. BUENOS AIRES: Dunes ca. 2 km SE of Argerich in "E p terrain of the Universidad del Sur, 37 km ur of Bahia Blanca, dtd 455*. 457*, * 465 (DUSS; 457 so C 455, 457 also M, MO). NEUQUEN: Sandy places in the E^ part of the city of Renan, ‘Santa 542* 558, 560, 569*, 575* (DUSS; 542, 569 also CTES, M; 542, 569, 575 also MO). Sandy and waste places in the irrigation ditches of the farm “Granja LU" in the SE ds of the city of Neuquén, Santarius 567*, 590*, 594, 595 (DUSS; 590 also CTES, M; 567, 590 also MO). Stony places at Río Limay, 6 lem E of Piedra del Águila, Santarius 597*. 598*. 604* (DUSS; 598 also CTES, M; 598, 604 also MO). oe specimens examined: TI! BUENOS AIREs: Necochea, Hicken 4662 (LIL). Laguna Brava near Gral. Pue dini ds 100 m, Descole in 1938 (LIL). San Clemente near Gral. Lavalle, Cabrera 4262 (GH). Bahía Blanca. Claraz in 1884 (G). Dunes near Bahia San Blas, Fabris & Schwabe 5012 (LP). Pigué near Saavedra, Burkart 4713 (MO). Sierra de Curamalal, Cabrera 5433 LP). Sierra La Tinta, El Sombrerito, 400 m, Spegazzini 41 (BAB). Est. Sta. Maria at Rio Colorado near Villarino, Hunziker 4452 (POM, SI). Campo “La Susana” near Est. Peralta, 320 m, Huidobro 1177 (LIL, NY, S, SD. Sierra de la Ventana, Molfino 46137 (BAB); Gomez 11785 (BAA, MO). LA PAMPA: Guatrache, Williamson in 1925 (SI). NEUQUEN: Bajada del Manzano, 20 km S of Zapala, Ancibor 90245 (BAA). Rio NEGRO: Choele Choel near Perella- neda, 152 m, O’Donell 1795 (NY). Dep. Conesa, China Muerte near Laguna del Mate in the valley of Rio Negro, 60 km E of Carmen de Patagones, Krapovickas 2078 (RSA); Hunziker 9 (CORD, RSA). cusur: Valle de Las Plumas, Gerling 137 (POM). Cabo Raso, Miiller 24 (CORD). Between Travesia de Rawson and the Seis Illin 16 (SI). It has not yet been possible to analyze the complexes involved in the origin of O. bahia-blancae with any degree of certainty. On the basis of the strigillose pubescence and the relatively small leaves, one might suggest that one of the complexes of O. mendocinensis is involved, while the other might be an altered complex derived from O. longiflora and with genes for smaller flowers than in that species. Such a complex, ultimately derived from O. longiflora, is repre- sented both in O. parodiana of series Allochroa and in O. cordobensis of series Clelandia. 550 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 25. Oenothera picensis pond Anales Mus. Nac. Chile, Bot. 1891: 22. 1891. Fics. 65-66, 140-142, 222 Annual herb with an erect or somewhat decumbent main stem, not forming a rosette, simple or branched near the base, 2-10 dm tall. Plants very densely to sparsely long-villous, the hairs soft, densely to sparsely short-villous and glan- dular-pubescent. Cauline leaves narrowly elliptic to lanceolate, acute, acute to truncate at the base, sessile, 3.5-10 cm long, 0.5-2 cm wide; bracts narrowly oblong or narrowly lanceolate to narrowly ovate, acute, truncate to subcordate at the base, sessile, 2.5-6 cm long, 0.5-2 cm wide, longer than the capsule they subtend or subequal to them, occasionally shorter; leaves plane or coarsely undu- late at the margins, irregularly serrate, the teeth blunt. Inflorescence mostly branched. Floral tube 1.54.5 cm long. Buds oblong to lanceolate in outline, 0.7-1.7 cm long, 3-5 mm thick, often red striped at the junction of the sepals with the floral tube. Sepals often flecked with dark red; apices of the sepals 1-2 mm long, mostly erect. Petals broadly obovate to very broadly obovate, sometimes broadly elliptic, 0.7-2.5 cm long. Anthers 5-12 mm long. Filaments 5-15 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 2-6 cm long. Stigma lobes 3-6 mm long. Ovary 1-2 cm long. Cap- sule (2-)2.5-4(-4.5) cm long, 3-4 mm thick. Seeds elliptic to broadly elliptic in outline, 1.2-2 mm long, 0.5-0.8 mm thick, brown. Self-pollinating complex heterozygote. Gametic chromosome number, n — 7 (ring of 14, ring of 10 and ring of 4 or ring of 8 and ring of 6 at meiotic metaphase I). Lectotype: Chile, Prov. Atacama, oasis of Pica, Mar. 1885, R. A. Philippi (SGO-52850, GH photograph). Distribution (Fig. 237): At the foot of the Andes in Argentina in the prov- inces of Jujuy, Salta, Tucumán, Catamarca, San Juan and Mendoza, and also in the provinces of Buenos Aires, Córdoba, San Luis, and Río Negro; on the western side of the Andes in Chile from the province of Antofagasta to Colcha- gua. Oenothera picensis subsp. picensis is adventive on the Juan Fernández Islands and in the provinces of Bío-Bío and Malleco in southern Chile. Each of the three subspecies of O. picensis is made up of chromosomal com- plexes originating from O. odorata on the one hand and O. affinis on the other. They must have originated independently from one another, however, judging from their completely distinct areas of distribution. One of the most important characteristics of O. picensis is that it forms no rosette, a characteristic that can be traced back to the influence of O. affinis, one of its parents. The soft pubes- cence is also reminiscent of O. affinis, whereas the width of the bracts at their base is a characteristic of O. odorata. KEY TO THE SUBSPECIES l. Plants densely E lor tube 2-4.5 cm long 25a. subsp. picensis Floral Ne l .5—2 cm long 25c. subsp. bonariensis l'. Pant jn 5 long -yillous, 1.5-2.5 cm long subsp. picensis F. Petals 0.7-1.3 cm long 778 poss cordobensis 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 551 40 | e ` í 4 S ac > ` 4 L mel) \ 8 | `" „ ae x j / J . ^ ur E.g { 47 0 . 7^; — | 800 KILOMETERS j j / j , j / / 30 20 iangles), 226. Ranges of Oenothera versicolor (circles), O. rubida (filled tr We eal ae 5 and O. ravenii subsp. chilensis (dots). iso 552 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 25a. Oenothera picensis subsp. picensis.—Fic. 140. odorata sensu Gillies ex Hooker, Bot. Misc. 3: 310. 1833, Pro parte. mollissima sensu Gillies ex Hooker, Bot. Misc. 3: 310. 1833. mollissima sensu Reiche, MEE Univ. Chile 98: 476. 1897. i err sensu Munz, Amer. J. Bot. 22: 659. 1935, pro parte; Revista Univ. (Santiago) : 266. , pro ds affinis sensu Munz, Re ath Univ. (Santiago) 22: 267. 1937, pro p dorata “Erlangen” Haustein, Z. Indukt. Abstammungs- UL MN 84. 418. 1952; Cle- "land. Jap. J. Genet. 43: 332. 1968. mollissima "Uspallata" Tandon & Hecht, Cytologia 21(3): 252. 1956. D. pe sao Plants 4-10 dm tall. Plants densely to sparsely long- and short-villous, sparsely glandular-pubescent. Cauline leaves narrowly lanceolate to lanceolate, acute, 6-10 cm long, 0.5-1.3 cm wide; bracts lanceolate to narrowly ovate, acute, truncate to subcordate at the base, 3.5-6 cm long, 0.8-1.5 cm wide. Floral tube 2-4.5 cm long. Buds oblong to lanceolate in outline, 1.2-1.7 cm long, 4-5 mm thick. Petals very broadly obovate, 1.5-2.5 cm long. Anthers 7-12 mm long. Filaments 11-15 mm long. Style 3-6 cm long. Stigma lobes 4-6 mm long. Ovary 1.2-2 cm long. Capsule 2.5-3.5 cm long. Seeds elliptic in outline, 1.3-2 mm long, 0.5-0.7 mm thick. Gametic chromosome number, n = 7 (ring of 14* at meiotic metaphase I). Flowering time: October-May. Distribution (Fig. 237): In Chile from the province of Antofagasta to Col- chagua; adventive on the Juan Fernández Islands and in the southern Chilean provinces of Bío-Bío and Malleco; on the east side of the Andes in the provinces of Mendoza and San Juan in Argentina. 5 examined from cultivated plants: VALPARAÍSO: Concón to Salinas near the border to Prov. Aconcagua, Hjertung & Rahn 553* (CTES, DUSS, M, MO). Concón, N of Viña del Mar, Gópel in 1961 (DUSS, MO). 1550* (DUSS; 1549 also CTES, M, MO). On rocks ca. 5 km S of Cacheuta, 23 km N of Luján de Cuyo, 1200 m, Santarius 1551, 1553 (DUSS; 1553 also MO). Stony and waste places ca. 2 km N of Tupungato, 1250 m, Santarius 1554*, 1556, 1564, 1565, 1570, 1571* (DUSS; 1554, 1564 also MO). At the road above E] Peral, 6 km NW of Tupungato, 1350 m, Santarius 1576* (DUSS, M, MO). ULTIVATED: O. “odorata” from the Botanical Garden in Erlangen, Germany, received d camined: CHILE. ANTOFAGASTA: 5 Martin nm E Toconao, 2,300 m, Ricardi 2998 (CONC); Ivanovic 7713 1 0 ATACAMA a, valley of Rio del Transito, 1,450 m, Johnston 5861 (GH, K, POM). Est. Manflas, eee 3748 (CONC). La Higuerita near San Félix, 1,300 m, Ricardi 10 (CONC). Copiapó, Ricardi 3625 ( CONC); Gigoux in 1885 (GH). Desert of Atacama, Morong 1105 (F, MO, NY). coquIMBo: enis Alamo 19175 (CONC). Punitaquí near Ovalle, Jiles 1936 (CONC), ACONCAGUA: rtillo on road to Mendoza, Diaz in 1861-62 (SGO). Rio Blanco, 1,400 m, Giinther & 1 ois in 1928 VALPARAÍSO: Bueck in 1843 (HBG); Wilkes (NY); Buchtien in 1895 (US). Olmué near Limache, N.N. 1901 (HBG). Limache, Behn 22798 (CONC). Reñaca valley, Poulsen in nN C). Günther & Buchtien in 1928 (HBG). santiaco: Claude-Joseph 726 (US). Romeral at Rio Yezo, Biese 83 (GH, LIL). Chacayes at Rio Yezo, Biese 139 (LIL). Dunes near El Tabo, Kohler 229 (CONC). San Antonio, Looser 924 (SI). Dunes of Llolléo S of San Antonio, Poulsen in Í Bra Valley x he m Obra and Canelo, Looser 144-17 (SI). El Manzano ne é de M 870 m, Montero in 1927 (GH, K); Sparre in 1947 (S). Batuco, Reiche gc 061514); "nes 9406 (POM), 257 (SI). corcHacua: Talcaregoné, 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 553 Gay in 1831 (P). Bío-Bío: Trapatrapa (SGO-052829). MALLECO: Sta. Rosa, agr in 1882 (SGO). Islas Juan Fernández: Philippi in i (SGO-052867 ); Bertero 1486 K, P). Masatierra, Biirger (GOET); Skottsberg 141 (S, UPS); Bock 22801 (CONC). Masafuera, Quebrada Casas, Meyer 9352 (MO). Province unknown: Aeuleo, Bertero 446 (W), 464, 1185 (G, MPU, NY, P). Cerro S. Cristóbal, N.N. in 1868 ( W). ARGENTINA. MENDOZA: Jörgensen 131 (C). Mendoza, 800 m, King 112 (BM); Sanzin 7 (SI); Loos 2390 (CORD). San Carlos, Roig 5178 (CORD); Covas 3492 (SI). Dep. J Campo de los Andes, 1,800 m, Paci 713, 727 (LIL); Barkley 20Mz126 (LIL, NY, L, NY). Leal 963 (Leal). Villavicencio, Burkart et al. in 1942 (SI-14176). Tupungato, Ruiz Leal 2784b, 2784 (Leal, POM), 2775 (Leal, LP). Dep. Lújan, Cacheuta, Ruiz Leal 8734 (Leal, LIL). Potrerillo, 1,500 m, Semper 401 (LIL); l E m, Garcia 374 (LIL). Dep. San Rafael, Río Diamante, Beales 1966 (LIL). Aguo del Sapo, 1,200 m, Ruiz Leal 7378 (Leal, LIL). Río Atuel, 1,400 m, Wilczek 410 (G). Between Log and Las Penas, Kurtz 5412 (CORD). Cordillera of Mendoza, Cordén de Sta. Elena, Figueroa 15736 (CORD). Cordilleras, Jensen- Haarup in 1904 (C, LD, S). Specimens from plants cultivated in garden eningrad, seeds from Chile sent by Tanka in 1833 (LE). Kew, in 1880 (K; as O. i2 ). Among the F, generation hybrids between O. affinis and O. picensis subsp. picensis there are two classes of plants, one closely similar to each parent. The same occurs in hybrids between O. odorata and O. picensis subsp. picensis. Consequently, the parentage of this particular subspecies seems to be clearly demonstrated. Since there is no evidence that chromosomally homozygous entities of this series have ever occurred in Chile, it seems most probable that the O. affinis complex in O. picensis subsp. picensis was derived from the complex heterozy- gous O. affinis that does occur in Chile, and that the O. odorata complex was derived via O. stricta. It is also possible, of course, that O. stricta has derived its odorata-complex from O. picensis. This seems improbable, however, since the ravenii-complex of O. stricta in Chile is much more widespread than the affinis- complex of O. picensis, so that one might surmise that the odorata-ravenii com- bination originated earlier than the odorata-affinis combination. From Chile O. picensis subsp. picensis seems to have migrated over the Uspallata Pass to Argentina, where it seems to form occasional secondary hy- brids with the homozygous O. odorata, judging by the presence of plants with relatively large flowers in the Argentine portion of the range of O. picensis subsp. picensis. 25b. Oenothera picensis subsp. cordobensis Dietrich, subsp. nov.—Fıcs. 65- O. mollissima sensu Munz, Physis 11: 282. 1933, pro parte. Oenothera parodiana sensu Munz, Physis 11: 283. 1933, pro parte; Amer. J. Bot. 22: 662. 1935, pro parte. Plantae 2.5-9 dm altae, plerumque pilis longis villosis dispersis praeditae, etiam pilis brevibus villosis glandulosibusque praeditae. Folia caulina anguste elliptica vel lanceolata, 3.5- 5 cm longa, 0.5-1 cm lata; bractea anguste oblonga ad oblongam vel anguste lanceolata at c qualia. Tubus dinalis 3-4.5 cm longus. Gemmae ambito oblongae vel lanceolatae, 0.7-1.2 cm longae, 3-4 mm crassae. Sepala fusco-rubro saepe punctata; apices sepalorum 1-2 mm longi. ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ° 100 200 300 400 500 600 MILES r M —— 0 200 400 eoo 800 MiLOMETERS 0 : N. N. (K). PAYSAN DU: Km 437 at Ruta 3 near ae Chapicuy, Millot 494 (MVFA). SORIANO: Est. 1 nA near Mercedes, N.N. 91 (POM) Wright in 1928 ( LonEs; Between Rio Yi and Arroyo Mariacho, 5 B1521 IL, POM). FLORIDA: Rosengurtt B703 (BAA). Mansavillagra, Rosengurtt B1657 (LIL, POM). CANELON Dunes near La Floresta, Steer in 1923 (HBG). MonTEvIDEO: Gibert 86a, 341 (K); Orbigny in 1829 (P). Quinta Narelo, Fruchard in is (P). Carrasco, Munz 15451 (NY, POM). CERRO LARGO: = ES Paso Cruz and Río Tacuari, Arrillaga 2341 (MVFA). Between ue Negro and Arroyo Aceguá, Bout 890 (POM). MALDONADO: Solís, dd 21653 (B REM); Cunningham in 1867 (K). ENTINA. FORMO Jorgensen 3025 (POM, SI). cHaco: Las Palmas, Jörgensen 2487 dde SL US), 2489 Mes p (POM). SANTA FÉ: Mocovi, Venturi 275 (SI). Chacras be- ween Carcarana and Canada de Ee Berndt 5166 ( CORD). Rafaelo near Castellanos, Terribile 387 (LIL). Misiones: Puerto Segundo near Iguazú, Montes 10363 (CORD, LP). Soberbio near Guarani, Crisci 297 (LP). conmigNrgEs: Gral. Paz, Krapovickas d» Cristóbal 11836 (CTES, UC); Schwarz 321 (LIL). Dep. Capital, Arroyo Riachuelo at Ruta 12, Kra- povickas & Cristóbal 13783 (BAA, BAB, C, CTES, LP, MO). La Cruz near San Martin, Parodi 12368 (BAA). Juan Pujol near Monte Caseros, Ibarrola 2323 (LIL). Est. Tranqueras near Monte Caseros, Nicora 5187 (CTES). Vicinity of 5 Ibarrola 022, 31, 546 (LIL). Vicinity of Paso de los uds. Schinini 7699 (C , MO); Huidobro 3815, `3837 (LIL). Est. Santa Teresa near Mburucuyá, Pedersen 93 pro Ros rte (C). Vicinity of Ituzaingó, Pierotti 6227, 6195, a A Tisi 2 Ibarrola 1438 (LIL, NY); Krapovickas et al 21150 (CTES, MO). cedes, Millan 334 (POM). Between Ruta 27 and Rio Corrientes near Esquina, Krapov ikas et al. 27111 trud MO). ENTRE RIOS: Moe ee Meyer 11187 (LIL). Colón, Meyer 10657 (LIL). Concordia, Burkart 26731, 28763 (MO); Cabrera 19261 (LP). iE del Uruguay, Lorentz 514 (BR EM, CORD, GOET, K, W). Est. La Selmira near Concordia, Pedersen 7260 (C). Federación, Crovetto 4889 (BAB). BUENOS AIRES: Est. San Juan, 30 km S of Buenos Aires, Eyerdam et al. 23037 (SI). Cajón, Cabrera 6430 (LP). Delta del Paraná, Arroyo Carabales, Burkart 4316 ( CTES). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 573 ° ioo 200 300 400 $00 — ni rt rt L 00 200 — 400 — $00 — MILES #00 KILOMETERS 50 40 30 20 FicurE 234. Ranges of Oenothera affinis (hollow triangles), O. villaricae (circles), O. brevipetala (dot), and O. featherstonei (filled triangles). 574 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 In habit, O. parodiana subsp. brasiliensis resembles the chromosomally ho- mozygous O. ravenii very closely. Analysis of the complexes present in this subspecies has shown that it is predominantly derived from O. ravenii and O. indecora, with some introgression from O. affinis and O. longiflora, as indicated among other points by the length of the floral tube. Transitional forms with subsp. parodiana occur frequently in the zone of contact. 29. Oenothera verrucosa Johnston, Contr. Gray Herb. 70: 77. 1924—Fics. 12, 149, 185, 209. Erect annual herb, not forming a rosette, simple or branched from the base upward, 1.5-5 dm tall. Plant moderately to sparsely strigillose, moderately to sparsely villous, and sparsely glandular-pubescent. Cauline leaves very narrowly elliptic to elliptic or lanceolate, acute, attenuate to rounded at the base, only the lowermost short-petiolate, the remainder sessile, 4-8 cm long, 0.5-1.5 cm wide; bracts narrowly lanceolate to lanceolate, acute, rounded to truncate at the base, sessile, 3-5 cm long, 0.5-1.2 cm wide, longer than the capsules they subtend; leaves plane to slightly undulate at the margins, remotely serrulate, the teeth blunt. Inflorescence simple or branched. Floral tube 0.6-1.1 cm long. Buds oblong to broadly elliptic in outline, yellowish, often flushed with red, 0.3-0.6 cm long, 2.5-3.5 mm thick; apices of the sepals 0.5-1 mm long, erect. Petals very broadly obovate, 0.3-1 cm long. Anthers 1-2.5 mm long. Filaments 2.5-4 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 0.8-1.5 cm long. Stigma lobes 1.5-2 mm long. Ovary 0.6-1 cm long. Capsule 1.5-2.5(-3) cm long, 2.5-3.5 mm thick, erect, apparently petiolate; valves spread- ing apart after dehiscence, not curving. Seeds elliptic in outline, 1.5-1.7 mm long, 0.7-0.8 mm thick, dark brown to almost black. Self-pollinating. Gametic chromosome number, n —7 (7 bivalents* at meiotic metaphase I). Flowering time: March-April. Type: Peru, Dep. Arequipa, ravines and rocky slopes, Pampa, southern slopes of Chachaní Mountain north of Arequipa, 3,660 m, Mar. 1920, Mr. and Mrs. F. E. Hinkley 17 (GH, holotype; BAS, isotype). Distribution ( Fig. 238): Known only from the Andes of the department of Arequipa, southern Peru, at elevations from 2,400-3,700 m. Specimens examined from cultivated plant Peru. Dep. Arequipa: Quebrada de San Láza , 5-6 km NNE of Arequipa below the El Misti, sandy places W of the river, 2,600 m, yet 2068*, 2071, 2073* (DUSS; 2068, 2073 also M; 2068 also CTES, MO). Additional ME examined: UIPA: Near Arequipa, e DIO (K); 2,700 m, N.N. in 1954 (RSA); 2, 400-2, 600 m, Pennell 13174 (F, GH, K, NY, S, US, USM); Nuñez 97 (USM); N.N. in 1925 (US-1231003). o de Jesus, 2,600-2, 700 m, Ferreyra 14252 (USM). Slopes of the Misti between Yura a eda. Sandeman 3960 (K). Mountains near Yura, 2,575-2,600 m, Vargas 7974 (LIL, MO, RSA). Quebrada San Lázaro, 2,800 m, Munz 15483, 15527, 15528 (POM). Chihuata 20 km E of Arequipa, 2,600 m, Munz 15540 ( POM, US). Characato near Arequipa, 2,400-2,600 m, Vargas 8043 (LIL). — d 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 575 Oenothera verrucosa seems to be a declining species which has found a last refuge in the Andes of Arequipa. It evidently participated in the origin of the derivative complex heterozygote O. arequipensis, which occurs at low elevations along the coast of southern Peru and northern Chile. Possibly O. verrucosa at one time occupied much of the same area as O. arequipensis, but more likely that derivative species was simply able to expand its range into the newly devel- oped very arid regions during and after the Pleistocene, whereas O. verrucosa became confined to its present very limited area of distribution. An important characteristic of O. verrucosa is the stipitate appearance of its erect capsules, owing to their abrupt narrowing toward the base. Oenothera verrucosa and the following two species, O. coquimbensis and O. arequipensis, are obligate annuals and do not form a rosette. Responding to the short period of growth available to them, they elongate rapidly after forming only a few leaves and form their first buds only 4-5 weeks after germination. In cultivation, all other species require a much longer time for the initiation of their first buds. 30. Oenothera coquimbensis Gay, Fl. Chil. 2: 331. 1847.—Fics. 150-151, 186. O. 1 Philippi, Linnaea 33: 68. 1864. Lecrorype: Chile, Prov. Atacama, near . 1854 pe P e (SGO-052835, GH, NY and POM photographs); Munz, An mer. T. Bot. 22: e O. E s esa fae ied e M (Philippi) Reiche, Anales Univ. Chile 98: 476. 1897; Chile: 258. Onothera albicaulis R var. tigrina H. Lév. subvar. Pig ls (Gay) H. Lév., Monogr. . 345. 1909; Bull. Acad. Int. Géogr. Bot. 05. Aaa coquimbensis (Gay) Sprague & Riley, as Misc. 1 1921: 200. 1921. Erect annual herb, not forming a rosette, simple or branched from the ground upward, 0.5-5 dm tall. Plants densely to moderately strigillose, moder- ately long- and short-villous. Cauline leaves very narrowly elliptic to elliptic or lanceolate to narrowly ovate, acute, acute to truncate at the base, sessile, 5-8 cm long, 0.5-1.5 cm wide; bracts narrowly lanceolate to narrowly ovate, acute, truncate to subcordate at the base, subsessile, 2-6 cm long, 0.5-1.5 cm wide; leaves usually plane at the margins, irregularly and coarsely toothed or + regu- larly and deeply toothed, sometimes doubly so, often flecked with dark red- dish brown. Inflorescence simple or branched. Floral tube 1-3 cm long. Buds broadly elliptic to narrowly ovate in outline, yellowish, often striped with red at the junction of the sepals with the floral tube, 0.5-1.3 cm long, 3-3.5 mm thick; apices of the sepals 1-3 mm long, erect or divergent. Petals very broadly ob- ovate, 0.8-2 cm long. Anthers 2.5-6 mm long. Filaments 5-13 mm long. Style held above the anthers in most individuals, occasionally shorter and then the anthers shedding directly on it at anthesis, 2-4 cm long. Stigma lobes 2.5-4 mm long. Ovary ca. 1 cm long. Capsule 1-2.5 cm long, 2.5-3.5 mm thick, + erect or spreading obliquely from the stem, often somewhat enveloped by the subtending ract. Seeds narrowly elliptic in outline, 1.2-1.6 mm long, 0.4-0.5 mm thick. Self-compatible but mostly outcrossing. Gametic chromosome number, n = 7 (7 bivalents* at meiotic metaphase I). Flowering time: September-November. Type: Chile, Prov. Coquimbo, on dunes at the edge of the sea, very rare, vicinity of La Serena, Dec. 1836, Cl. Gay 520 (P, holotype; F, GH, K, P, isotypes). 576 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Distribution (Fig. 239): Sandy flats and dunes near the coast in the semi- deserts of Chile from Antofagasta to Purén, Malleco Province. eng ud = from cultivated plants: Grown from seeds of a herbarium specimen in 1973 at MO. Source: Chile, Prov. Ata- cama, Dep. Freirina, road from Chañaral de Aceituna to Bahía Carrizal at km 6, Marticorena, „ Weldt 1846* ( CONC-36723). Additional specimens examined: Cu Base of hills just SE of La Chimba, Johnston 3640 (GH, US). ATACAMA: Geisse in 1885-1887 (NY); Morong 1141, 1162, 1231, 1264 (NY), 1285 (K, MICH, NY, US). Chanaral, Ricardi 2258 (CONC). Hills back of El Barquito at Puerto de Chañara l, Johnston 4772 (CH). Vicinity of Caleta Pan de Azúcar, dunes on point just S of the Caleta, Johnston 5839 (F, GH, K, LIL, S, US). Road from Chanaral de Aceituna to Bahia Car- rizal at km 6, Marticorena 1846 (CON C), Carrizal, N.N. in 1885 (SGO-052891). Huasco, Monypenny 46 (CONC). Algarrobal at Pan-Amer. a Cabrera 12649 (LP). Ricardi 4410- C). I km S of Huasco, Bócher et al. 540 (C). Vicinity of Copiapó, 370 m, Johnston 4990, 5030 (GH); Philippi 1726 (SGO); N.N. in 1885 (SGO-052892). Pan-Amer. Highway between Caldera and Chañaral at km 18, Ricardi et al. 1304-B (CONC). Between Copiapó and Vallenar at km 55, Ricardi et al. 1511 (CONC). At km 40, Ricardi 2210 (CONC). Morro de Copiapó, sandy washes along a stream near the sea, Worth 16185 (K, UC). Quebrada de Chanchoquin near Copiapó, Gigoux in 1895 (GH). Travesia, Kohler 153 (CONC); Jiles 2160 C); N.N. (SGO-052890). Caldera near Copiapó, Gigoux in 1894 (GH); Pod 1726b, 1726c, 1724 (SGO), in 1885 (SGO-052889). Piedra Colgada, Philippi in 1885 (SGO-052893, SGO-052887). Bandumias, Geisse 1726a (SGO). Pojonales, Geisse in 1888 (SGO). co- QUIMBO: Gay in 1838 (P); Reiche in 92 0 (SGO-052879); Jaffuel 2679 (GH); Philippi 785 (US); Biot 85 (K). Between La Serena and Punta de Teatinos, West 3919 (GH, MO, UC) La Serena, Choros Bajos, Marticorena 1694 (CONC). Between Herradura and Coquimbo, Skottsbe rg in 1917 (NY, S). Dunes between Tongoy and 5 near Ovalle, Gleisner 5 (CONC). Est. Talca near the sea, dep. Ovalle, Tiles 1425 (C S). Quilimari, in 1862 (SGO-052833). vaALPARAÍsO: Valparaiso, Calvert in 1914 "EMT "MALLECO: Near Purén, N.N. 1229 in 1838 (SGO-052834 ). Cultivated specimens: Botanical Garden, Leningrad, seeds from Chile, Prov. Atacama, from sandy plains near Huasco, sent by Cuming, in 1847 (LE; as O. glauco-virens F.M.). Oenothera coquimbensis is totally distinct from O. verrucosa in its character- istic pattern of toothing of the leaves, its larger flowers, and its different mode of capsular dehiscence, in which the valves curve inward. In addition, the cap- sules usually spread obliquely from the stems instead of standing erect and are often partially enfolded by their subtending bract. Finally, the seeds are shorter and narrower than those of O. verrucosa. Within O. coquimbensis there are two distinctive variants, but these are so completely joined by intermediate forms that it does not seem desirable to accord them formal taxonomic recognition. One has extraordinarily deeply toothed leaves in which the teeth are very narrow and long and often also sec- ondarily toothed. The second has relatively wide leaves which are merely coarsely serrate. Oenothera coquimbensis, O. arequipensis, and to some extent, O. nocturna, species of the coastal deserts of Peru and Chile, appear only in years of ample rainfall, and thus represent a distinctive ecological type within subsect. Munzia. The evolution of these deserts, and consequently of the species that inhabit them, is a phenomenon of Late Pleistocene and Recent time (Raven & Axelrod, 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 577 31. Oenothera arequipensis Munz & Johnston, Contr. Gray Herb. 75: 20. 1925. —Fics. 152, 187-188. O. laciniata Hill var. limensis Munz & Johnston, Contr. Gray Herb. 75: 20. 1925. Type: Per Dep. Lima, sandy lomas along the sea near Lurin, 23 Sep. 1925, J. F. Macbride 5950 (F-536954, holotype, NY, photograph; GH, K, US isotypes). O. laciniata var. nocturna (Jacq.) Munz, Amer. J. Bot. 22: 656. 1935, pro part O. laciniata sensu Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(4): 537. 1941. pro parte. O. verrucosa sensu Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(4): 540. 1941, pro parte. Erect annual herb, not forming a rosette, usually branched near the base, 1-3.5 dm tall. Plant densely to sparsely strigillose and densely to sparsely ap- pressed- or erect-villous, sometimes almost glabrous. Cauline leaves narrowly elliptic to narrowly oblanceolate, acute, gradually narrowed to the short petiole, 2-10 cm long, 0.5-2 cm wide; bracts narrowly elliptic to narrowly ovate, acute, acute at the base, sessile, 2.5-5 cm long, 0.5-1.5 cm wide, longer than the capsules they subtend; leaves plane to slightly undulate at the margins, usually deeply sinuate. Inflorescence usually branched. Floral tube 1-3 cm long. Buds oblong in outline, yellowish, sometimes flushed with red, 0.3-0.8 cm long, 2-4 mm thick; apices of the sepals 1-2 mm long, erect or divergent. Petals very broadly obovate, 0.4-1.5 cm long. Anthers 2-3.5 mm long. Filaments 5-7 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 1.3-4 cm long. Stigma lobes 2-3.5 mm long. Ovary ca. 1 cm long. Capsule 1.5- 3 cm long, 2.5-4 mm thick, tapering at both ends and usually appearing pedicel- late, mostly erect; valves spreading apart in dehiscence. Seeds elliptic to nearly rotund in outline, 1-1.3 mm long, 0.6-0.8 mm thick. Self-pollinating complex heterozygote. Gametic chromosome number, n = 7 (ring of 14* at meiotic metaphase I). Flowering time: September-November. Type: Peru, Dep. Arequipa, sandy slope, desert hills near Mollendo, 17 Nov. 1923, A. S. Hitchcock 22403 (US-1,196.655 ). Distribution (Fig. 231): In the lomas of the Pacific coastal deserts and semi- deserts of Peru from the department of Libertad to Tacna, ascending to 2,700 m elevation in the vicinity of Lima; in Chile only known from Tocopilla in the province of Antofagasta. Specimen ee from cultivated plan Peru. tra: Prov. Huarochiri, Huascamarca near Santiago de Tuna, 2,700 m, Encarna- ción 346* (DUSS Additional specimens examined: PERU. LA LIBERTAD: Cerro Campana near Trujillo, 350 m, López 0907 (US). Cerro Ca- bezon near Trujillo, 400-500 m, Weberbauer in 1940 (USM). ANCASH: Lomas de Casma near Santa, 250-300 m, Ferreyra 8039 (US). Lima: Lima, Cuming 1079 (K); Soukup 2158 (F); Mathetos pro parte (K). Lomas near Barranco, 50-100 m, Weberbauer 5703 (F, US). San Agustin, Asplund in 1940 (RSA). Lomas dé Atocongo, 300—400 m, Ferreyra 2448 US). Lomas de Lachay near Chancay, Cerrata 3825 (MO). Pasomayo near Chancay, 300 Stork 9353 (GH, K, UC). Hacienda Desagravio near Huaura, més dan in 1943 (USM- 12875). AREQUIPA: Arequipa, in 1892 (GH). Mollendo near Islay, 300 m, Ferreyra 12102 (USM); Mexia 04167 (MO, UC, US); Stafford 279 (K). Hillside dde Duck of the port, Johnston 3556, 6303 (GH); Worth & Morrison 15764 (GH, K, UC). Slopes of the Misti, Cárdenas & Rodrígu iez 3 (F). Prov. Caraveli, Lomas de Jahuay, km 534 between Nazca and Chala, 300-400 m, Ferreyra 1396 (USM). Lomas de Capac near Chala, 300 m, Ferreyra 1433 (USM). Between Tambo and Posco, 300-550 m, princess Therese of Bavaria 233 (M). Lomas de Camaná, 600-700 m, Ferreyra 6440 (US, USM). Cerro los Cerillos, W of Pan Am. High- Fic (hollow ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 f l n * e fal NA io ë A f N ( URE 235. Ranges of Oenothera stricta subsp. argentinae (circles), O. montevidensis triangles), O. pseudolongiflora (filled triangle), and O. hechtii (dot). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 579 way, 52 km S of Nazca, km 500, 650 m, Rahn 130 (C). Along Pan-Amer. Highway, 28.9 km NW T 240 m, Rahn in 1957 (C). Capac, Scolnik 1019 (CORD, LIL, RSA). MoguE- cua: Lom s de Ilo near Mariscal Nieto, 600-700 m, Ferreyra 12577 (BM, MO, USM). Lomas Mostacilla near Ilo, 300 m, Vargas 8598 (LIL). TACNA; Morro de Sama, 450 m, Delarte CHILE. ANTOFAGAST Tocopilla, steep hillside ca. 6 km north of port and about opposite Caleta Duendes, g 3602 (GH, S, US). Tocopilla, Jaffuel 2554 (GH). The close similarity in general aspect between O. arequipensis and O. verru- cosa led Munz (1935) to combine them as a single species ten years after first describing the former. Notwithstanding this, there are fundamental differences between them. Oenothera arequipensis is branched to a greater extent and has sinuate leaves and seeds that are much broader. Moreover, O. verrucosa is chro- mosomally homozygous and occurs only at elevations greater than 2,000 m in the department of Arequipa; O. arequipensis is a complex heterozygote and inhabits mainly the sea-level lomas of the coastal deserts of Peru and northern- most Chile. No hybrids involving O. arequipensis have yet been analyzed to determine the genomes involved in the formation of this species. Nevertheless, it probably contains a genome derived from O. verrucosa, since the resemblance between these two species is much too close to be attributed to chance. Its second ge- nome is probably derived from O. laciniata subsp. pubescens, which ranges south at least to the Peruvian Departments of Lima and Junin. At the very least, the second genome of O. arequipensis must contain a strong admixture of genes from this entity, as indicated by the sinuate leaves and the seeds which are often nearly rotund in outline. On the other hand, O. arequipensis never exhibits the nodding buds of O. laciniata. Oenothera arequipensis provides an impressive example of the reticulate relationships between the subsections of sect. Oenothera, linking a highly de- rived South American species with a “late arrival” in South America as a com- plex heterozygote of rather local distribution in a habitat that is clearly marginal for the genus as a whole. 32. Oenothera grisea Dietrich, sp. nov.—Fics. 86-87, 154. erba annua erecta, non rosulata, eae a, 2-5 dm alta, nonnisi denseque strigulosa, L 7 foliisque ut videtur erii ibus. Folia caulina anguste elliptica vel anguste lance- olata, acuta, basi acuta vel r ata, sessilia, 5-10 cm longa, 0.8-1.2 cm lata; bractea lanceolata vel ovata, acuta, basi wana vel subcordata; sessilia, 2-3 cm longa, 1-1.5 cm lata; folia manifeste ad margines undulata, irregulariter obtuseque serrata. Inflorescentia ramosa. Tubus floralis 1-1.5 cm longus. Gemmae ambito oblongae vel ellipticae, 0.8-1 cm longae, 3-4 mm crassae, griseo-virides; apices sepalorum 1-1.5 mm longi, divergentes. Petala latissime obovata, 0.8-1.2 cm longa. Stylus brevis, stigmate sub anthesi antheris circumdato. i 6-7 mm ee oe 2-2.5 cm longa, 2.5-3.5 cm crassa. Semina ambito elliptica, 1.3-1.5 mm lon 0.5-0.7 m assa. PURSE gameticus chromosomaticus, n — 7; planta chromo- L. 5 phos Erect annual herb, not 9 a rosette, usually with many branches arising from the base upward, 2-5 dm tall. Plants exclusively and densely strigillose, the stems and leaves appearing gray green. Cauline leaves narrowly elliptic to nar- rowly lanceolate, acute, acute to rounded at the base, sessile, 5-10 cm long, 0.8-1.2 cm wide; bracts lanceolate to ovate, acute, truncate to subcordate at the 580 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 base, sessile, 2-3 cm long, 1-1.5 cm wide; leaves evidently undulate along the margins, irregularly serrate with blunt teeth. Inflorescence branched. Floral tube 1-1.5 cm long. Buds oblong to elliptic in outline, 0.8-1 cm long, 3-4 mm thick, gray green; apices of the sepals 1-1.5 mm long, divergent. Petals very broadly obovate, 0.8-1.2 cm long. Anthers ca. 4 mm long. Filaments 5-6 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 1.5-2.5 cm long. Stigma lobes ca. 3 mm long. Ovary 6-7 mm long. Capsule 2- 2.5 cm long, 2.5-3.5 cm thick. Seeds elliptic in outline, 1.3-1.5 mm long, 0.5-0.7 mm thick, brown. Self-pollinating complex heterozygote. Gametic chromosome number, n — 7 (ring of 14* at meiotic metaphase I). Flowering time: December- March. Type: Grown from seeds and cultivated in the Botanical Garden of Düssel- dorf, Germany, 17 Aug. 1971. Source: Chile, Prov. Valparaíso, Las Ventanas, end of 1965, L. Constance ( MO-2155203, holotype; DUSS, M, isotypes). Distribution (Fig. 232): Known only from the dunes at Concón, province of Valparaíso, Chile. Specimens examined from cultivated plan CHILE. VALPARAISO: Las Ventanas, PLUS in 1965* (DUSS, M, MO). Additional specimens examined: HILE. VALPARAISO: Ritoque, dunes near Concón, Poulson in 1952 (C). Quinteros near Concón, Philippi in 1866 (W). Dunes near Concón, Jaffuel 3956 (GH); Zöllner 6086 (L). The complex heterozygote O. grisea is similar in habit to O. coquimbensis, but can be distinguished by its exclusively strigillose pubescence; plants of O. grisea are grayish in appearance. This species grows poorly in cultivation, and the flowers often drop off prematurely or do not form any pollen. For this reason, it has not yet been possible to analyze its genetic constitution fully. However, one genome seems to have been derived from the odorata-complex of O. stricta because hybridization with O. odorata produces an F, generation in which one type is very similar to O. odorata. The second genome may have been derived from the chromosomally homozygous O. coquimbensis, as sug- gested by the morphological similarity between that species and O. grisea. Both of the putative parents, O. coquimbensis and O. stricta, grow in the same area as O. grisea. The restricted distribution of O. grisea, which occurs only on the dunes at Concón near Valparaíso, seems to indicate a very recent origin for this species. 33. Oenothera featherstonei Munz & Johnston, Contr. Gray Herb. 75: 19. 1925. —Fics. 153, 189. Erect annual or perhaps sometimes perennial herb with arcuate side branches, to 5 dm tall. Plants exclusively strigillose. Cauline leaves very nar- rowly elliptic to elliptic, acute, narrowly cuneate at the base, short-petiolate, 3-5 dm long, 0.8-1.2 cm wide; bracts narrowly elliptic to lanceolate, acute, at- tenuate at the base, 2.5-4 cm long, 0.5-1 cm wide; leaves plane at the margins, irregularly serrate, the teeth sharp or blunt. Inflorescence branched. Floral tube 3-4 cm long. Buds lanceolate in outline, 2-3 cm long, 5-9 mm thick, yellowish; 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 581 | 0 100 200 300 400 30 $90MILES o 200 400 #00 800 KILOMETERS. ` 90 ‘80 50 40 I 30 20 Ficure 236. Ranges of Oenothera bahia-blancae (filled triangles), O. parodiana subsp. brasiliensis (circles), O. elongata (dots), and O. tucumanensis (hollow triangles). 582 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 apices of the sepals 2-4 mm long, divergent. Petals very broadly obovate, 2.5-4 cm long. Anthers 8-12 mm long. Filaments 16-24 mm long. Style long, the stigma elevated well above the anthers at anthesis, 4-7.5 cm long. Stigma lobes 3-7 mm long. Ovary 1.5-2 cm long. Capsule 2-2.5 cm long, 2.5-3 mm thick. Seeds elliptic in outline, 1.3-1.5 mm long, 0.6-0.7 mm thick, dark brown to almost black. Self-compatible but outcrossing. Gametic chromosome number, n — 7 (7 bivalents*, ring of 8 and 3 bivalents** or ring of 10 and 2 bivalents*** at meiotic metaphase I). Flowering time: February-May. Type: Peru, Dep. Lima, sprawling on disintegrated granite slope, Matucana, 12 Apr.-3 May 1922, J. F. Macbride & Featherstone 270 (F-516,803, NY photo- graph; G, GH, K, isotypes). Distribution (Fig. 234): Apparently only in the vicinity of Matucana, depart- ment of Lima, Peru, at elevations of 2,000-2,500 m. Specimens examined from cultivated plants: ERU. LIMA: Prov. Huarochiri, at km 63 of the railroad from Lima to Oroya, a een Surco and Puente Quitasombrero, 2, 050-2, 100 m, Encarnación in 1974*, **, *** (DUSS). 0 ional specimens examine LIMA: At Lima-Oroya railway between Surco and Matucana, 2,000-2,400 m, ELS 5217 (F, G, GH, POM, US). Matucana, 2,400 m, Asplund 10994 (RSA, UPS); Rauh in 1954 (RSA ). Vicinity of Surco, Ferreyra 9657 (USM). Puruchuca near Matucana, Mathews 492 (K). Near Canta, 2.200-2 500 m, Acleto 605 (USM). N.N. in 1950 (USM). Without exact locality: Martinet in 1878 (P), Martinet 71 (P). Oenothera featherstonei has flowers as large as those of O. odorata and O. ravenii. Its area of distribution presumably extended to lower elevations along the coast of Peru during the Pleistocene. Oenothera nocturna, its complex het- erozygote derivative, still coexists in these areas with O. laciniata subsp. pubes- cens. The spread of O. nocturna may have played a role in limiting the range of O. featherstonei to its present very limited area. Plants with 8 or 10 rings of chromosomes at meiotic metaphase I may represent hybrids between O. feather- stonei and O. nocturna. On the basis of its nearly black seeds and strigillose pubescence, which gives the plants a grayish hue, one might hypothesize that O. featherstonei may have been derived from an ancestral form similar to € peruana (series Renneria), the least specialized of all species of subsect. Munzia. 34. Oenothera nocturna Jacq., Coll. 3: 205. 1789; Icon. Pl. Rar. 3: 3, tab. 455. 1791.—Fi:cs. 155, 190, 224. O. albicans Lam., Lad Méth. 4: 552, tab. 279, fig. 2. 1797. TYPE: not seen. The illustra- tion in O. prostrata Ruiz & Pavón, Fl. Peruv. Chil. 3: 79, tab. 315. 1809. LECTOTYPE: Peru, Dep. Lima, common in the provinces of Lima and Chancay, 7 n (P). Onagra nocturna pe, Moench, Suppl. Meth. Pl. 287 ji e polymorpha H. Lév. race longiflora (Jacq.) H. T var. nocturna (Jacq.) H. Lév., mogr. Onoth. 364. 1909; Bull. Acad. Int. Géogr. Bot. 19: . 1909. Sus nocturna (Jacq.) Sprague & Riley, Kew Bull. 1921: 201. 1921. Oenothera laciniata Hill var. nocturna (Jacq.) Munz, Amer. J. Bot. 22: 656. 1935. O. laciniata sensu Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(4): 537. 1941, pro parte. = — Erect annual to perhaps sometimes perennial herb, forming a weak rosette, well branched near the base, 3-6 dm tall. Plants exclusively strigillose. Cauline 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 583 leaves elliptic or narrowly lanceolate to lanceolate, acute, narrowly cuneate at the base, short-petiolate, 4-6 cm long, 0.5-1.5 cm wide; bracts narrowly elliptic to lanceolate, acute, acute at the base, 1.5-3 cm long, 0.5-1 cm wide; leaves plane to slightly undulate at the margins, + regularly sinuate. Inflorescence usually branched. Floral tube 1.5-2.5 cm long. Buds oblong in outline, 0.5-1 cm long, 3-4 mm thick; apices of the sepals 1.5-2.5 mm long, divergent. Petals very broadly obovate, 1-1.5 cm long. Anthers 4-7 mm long. Filaments 6-13 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 2-3.2 cm long. Stigma lobes 3-5 mm long. Ovary 1-1.3 cm long. Capsule 1.8- 2.5(-3) cm long, 2.5-3.5 mm thick. Seeds elliptic to broadly elliptic in outline, 1.3-1.8 mm long, 0.8-0.9 mm thick, dark brown to almost black. Self-pollinating complex heterozygote. Gametic chromosome number, n=7 (ring of 14* at meiotic metaphase I). Flowering time: At high elevations, February—June; at low elevations, September-November. Neotype: Jacq., Icon. Pl. Rar. 3: tab. 455. 1795. No specimen collected be- fore 1792 seems to have persisted, but the identity of this taxon is made clear by the plate selected as the neotype, and by the following two old cultivated speci- mens labeled with this name: Hort. Kew, 1792 (BM). Cult. Hort. Paris, Oct. 1815, Herb. J. Gay (GH, K). It was said to be from the Cape of Good Hope, but no African material has been seen; the original seeds undoubtedly came from the vicinity of Lima or elsewhere in Peru, to which the species is endemic. Distribution (Fig. 239): Predominantly at lower elevations in the depart- ments of La Libertad, Lima, Huancavelica, and Ancash, Peru, but ascending into the mountains along river valleys to 3,200 m elevation. Specimens examined from cultivated plants ERU. LIMA: Valley of Río E 15 Ou oad and on dry slopes, between the road (km 74.9-76.1) and the d ( km 00) from Lima to Oroya, near Matucana, ca. 2,200 m, Santarius 2327*, 2328*, 2333* 9 MO: 2328 also CTES, M). Additional specimens examine ERU. LA LIBERTAD: In valley between Pacasmayo and railhead, 2,130 m, Forbes in 1912 (BM). Trujillo, Barranza, 60 m, Sagdstegui 7857 (CTES, MO). AN CASH: Recuay near Marca, 2,600 m, Gómez 38 (USM). Carancayo in valley F Sipe near Bolgnesi, 2,600 m, Cerrate 12188 (BM, USM). Taclán, 3,050-3,100 m, Proaño 87 (USM). Lima: Lima, Savatier 1395 ( Martinet 40 (882) (P, US); Andeak in 1852 (S); EL 493 (K). San Agustin, Asplund 13824 (RSA). San Boskobo, 120-240 m, Saunders 163 (BM). Lomas de Lurín, 400— 500 m, Ferreyra 9536 (BM, USM). Lomas de Atocongo, 300—400 m, Ferreyra 0172, 2062, 12478 (USM); Aguilar in 1948 ( USM); Pennell 14777 (F); Vargas et al. 9296 (GH, K, UC). Chancay, Dombey 727 (G, L, P). Vargas 4710 (MO). Ruins of Cajamarquilla near Chancay. 300-400 m, Ferreyra 2838 (USM). Supe, near Chancay, 100 m, Goodspeed et al. 17361 (UC). Lomas de Lachay, between Chancay and Huacho, 400-500 m, Ferreyra 8770, 11503 (USM); Infantes 2128 (LIL); Cerrate 856 (BM, USM). Madalena near Lima, Née (F). San Isidro, Raimondi 6115 (USM). Miraflores, Maisch 13727 (USM). T do Jaffuel 3934 (GH). River Rimac, 1,220 m, Safford in 1887 (NY); Ball 1882 (G o parte, K, NY). Chosica, Martinet 40 (P, RSA). Huaquicha near Surco, 2,600-2 E m, Qm 6067 (US, USM). Surco, 2,000 m, Asplund 11056 (RSA). Santa Eulalia valley near Huarochiri, Mc. Hanrish 13, . Matoa, Raimondi in 1877 (USM). Mountains E of Tupe, Atsmito, 3,100 m, Cerrate 1059 (BM). Canta, 3,000-3,200 m, Pennell 14603 (F, GH). Cajatambo, 2,740 m, Sandeman 5322 (K). HUANCAVELICA: Córdoba near Castrovirreina, 3,050-3,300 m, Metcalf 30288 (G, GH, MO, UC, US). ns: en, Germany, 1795, Herb. Schreber 159 (M; as O. capensis). Erlangen, in 1800 bo. prn ni a Gottingen, Germany, in 1803 (LE). [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN !00 200 %% 400 500 4oowitts [LJ 400 KLOMETERS L " v — 4 20 60 Ficure 237. Ranges of Oenothera picensis subsp. picensis (hollow triangles), subsp. cordobensis (circles), and O. picensis subsp. bonariensis (filled triangles ). O. picensis 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 585 The complex heterozygote O. nocturna may be separated from the bivalent- forming O. featherstonei by its smaller flowers, autogamous breeding system, and usually sinuate leaves. The overwhelming morphological similarity be- tween these species leaves no doubt that O. featherstonei contributed one of the chromosomal complexes to O. nocturna. The pitting of the seeds in O. nocturna (Fig. 190) is very similar to that in O. laciniata subsp. pubescens (subsect. Rai- mannia). It seems possible that O. laciniata subsp. pubescens has contributed a genome to O. nocturna, and perhaps also to O. arequipensis (the other parent being O. verrucosa), but these hypotheses have not yet been tested experimen- tally. Neither O. nocturna nor O. arequipensis has the nodding buds character- istic of O. laciniata subsp. pubescens. Series III. CLELANDIA Oenothera sect. Oenothera subsect. Munzia series Clelandia Dietrich, ser. nov. Raimannia sensu Sprague & Riley, Bull. Misc. Infor. 1921: 200. 1921, pro parte. Oenothera 5 sensu Munz & Johnston, Contr. Gray Herb. 75: 16. 1925, pro parte. Oenothera subgen. Raimannia sensu Munz, Physis 11: 279. 1933, pro parte; Amer. J. Bot. 22 645. 1935, pro ak ~ erbae annuae vel biennes (O. punae perennis est), erectae vel prostratae, rosulatae vel erosulatae. Capsula sursum gradatim angustata, specieribus paucis cylindricis, + erecta, mani- feste bractea subtenta connata; valvulae capsulae post dehiscentiam extro curvatae. Annual or biennial herbs (only Oenothera punae perennial), erect or pros- trate, forming a rosette or the stem elongating soon after germination, un- ranched or with oblique or arching side branches arising from the rosette; plants 2-15(-20) dm tall, or with prostrate branches 5-25 cm long. Stems usu- ally thicker than those of series Allochroa, (3-)5-15 mm thick. Plants (1) densely to sparsely strigillose and densely to sparsely long- and short-villous; (2) densely to sparsely strigillose, densely to sparsely long- and short-villous, and moderately to sparsely glandular-pubescent; or (3) densely to sparsely long- and short-villous and densely to sparsely glandular-pubescent. Rosette leaves linear to narrowly oblong, very narrowly elliptic to elliptic or narrowly oblanceolate, gradually narrowed to the petiole or subsessile and narrowly cune- ate at the base, 2-25 cm long, 0.1-2.5 cm wide; cauline leaves linear to narrowly oblong, very narrowly elliptic to elliptic or narrowly lanceolate to lanceolate, acute, narrowly cuneate to truncate at the base, sessile or short-petiolate, 1.5-20 cm long, 0.1-2 cm wide; bracts linear, very narrowly elliptic to narrowly elliptic or narrowly lanceolate to narrowly ovate, acute, acute to subcordate at the base, 1.5-6 cm long, 0.1-1.5 cm wide, often with red margins; leaves plane or evidently undulate at the margins, usually irregularly serrate with blunt or sharp teeth. Inflorescence simple or branched; flowers erect or somewhat oblique with respect to the stem. Floral tube 0.5-10 cm long. Buds narrowly oblong to oblong, elliptic to broadly elliptic, or narrowly lanceolate to lanceo- late in outline, green to yellowish, often flushed with red, often with red stripes at the junction of the sepals with the floral tube, 0.3-2.5 cm long, 2-6 mm thick. Sepals rarely flecked with dark red; apices of the sepals 0.5-3 mm long, erect 586 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 | | | e | * 60 so 40 30 20 FicurE 238. Ranges of Oenothera parodiana subsp. parodiana (hollow triangles), O. punae (dots), O. verrucosa (filled triangles), and O. laciniata subsp. pubescens (circles). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 587 or divergent. Petals obovate to very broadly obovate, 0.4-3 cm long, yellow, rarely with an indistinct red spot at the base of each one. Style short, the anthers shedding pollen directly on the stigma at anthesis. Ovary 0.5-2 cm long. Cap- sule 1.24 cm long, 2.5-5 mm thick, in most species gradually narrower upward from a broad base, in a few broadly cylindrical, + erect, evidently fused with the subtending bract; valves curving outward as in series Allochroa when the capsule sheds its seeds. Seeds elliptic to rotund in outline, 0.8-2 mm long, 0.5- 0.9 mm thick, light to dark brown, sometimes flecked with dark red-brown spots. Self-pollinating complex heterozygotes. Gametic chromosome number, n = 7 (ring of 14 at meiotic metaphase I; plants with smaller rings very rare). Type species: Oenothera elongata Rusby. Distribution (Fig. 7): Most species of this series occur at low elevations. Only the species that occur in Bolivia and O. punae are characteristic of the high mountains. This series is dedicated to the late Ralph E. Cleland (1892-1971), student of Oenothera. All of the species assigned to it combine one genome derived from series Renneria with another derived from series Allochroa. See also the remarks on pp. 427 and 434 concerning the relationships of this group. Many of the species included here are relatively difficult to recognize as members of series Clelandia in a pressed condition. The capsules are mostly not cylindrical, however, and they do taper toward the apex. Whereas the cap- sules of series Allochroa always stand out obliquely from the stem, those of series Clelandia are more nearly erect, like those of series Renneria. In complex heterozygotes that involve O. affinis, however—at least in O. elongata and O. pseudoelongata—the capsules are cylindrical, which might be related to the fact that the capsules in O. affinis are somewhat swollen in their upper third. The inflorescence of series Clelandia is, as a rule, thicker and more heavyset than that in series Allochroa. 35. Oenothera magellanica Philippi, Anales Univ. Chile 84: 633. 1893.—F'cs. 83, 156 O. bo ayy Meigen, Bot. Jahrb. Syst. 17: 260, 291. 1893; non mu Steud., Nom. Bot., ed. 2. 206. 18 LECTOTYPE: Chile, Yerba Loca, 2.000 1 , 7 Aug. 1892, F. Meigen 539 GH Dae ), O. e Philippi var. 5 Macloskie, Rep. Princeton Univ. Exped. Patagonia 8 , 3): 613. 1905. TYPE: not loc Oenothera stricta sensu Macloskie, Rep. pU 5 0 Exped. Patagonia 865, 3): 614. 1905; sensu pon Physis 11: 284. 1933, pro parte; Amer. J. Bot. 22: 661. 1935, pro parte; sensu Bgche Onothera Such: 1 . Lév. race odorata (Jacq.) H. Lév. var. Pu cum op nile) H. Lév., Monogr. Onoth. 363. 1909; Bull. Acad. Int. Géogr. Bot. 19: Oenothera mollissima sensu Munz, Physis 11: 282. 1933, pro parte; sensu 55 Dansk. Bot. Ark. 22: 968. Oenothera odorata sensu Munz, Physis 11: 284. 1933, pro parte; Amer. J. Bot. 22: 660. 1935, o parte; Revista Univ. (Santiago) 22: 264. 1937, pro parte Erect annual or biennial herb, forming a rosette, unbranched or with a branched main stem and arching or obliquely ascending side branches arising 588 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 e 100 — 200 300 400 soo «soo MNES | — - i | ° 200 400 200 wiLOMETERS . PEN li 30 40 30 20 239. Ranges of Oenothera parodiana subsp. strigulosa (circles), O. acuticarpa (filled D. O. coquimbensis (hollow triangles), and O. nocturna (dots). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 589 from the rosette, 2-12 cm tall. Plants densely to very sparsely strigillose, mod- erately to sparsely long- and short-villous, rarely sparsely glandular-pubescent. Rosette leaves linear to narrowly oblanceolate, acute, gradually narrowed to the petiole, 10-25 cm long, 0.9-1.5 cm wide; cauline leaves very narrowly elliptic to narrowly lanceolate or narrowly oblanceolate, acute, narrowly cuneate to attenu- ate at the base, sessile, 5-20 cm long, 0.5-1.2 cm wide; bracts narrowly lanceolate to lanceolate, acute, rounded to truncate at the base, sessile, longer than the capsules they subtend, 4-6 cm long, 0.5-1 cm wide; leaves plane or markedly undulate at the margins, regularly or irregularly serrate, the teeth blunt or sharp. Inflorescence simple or branched. Floral tube 1.3-2.5(-3) cm long. Buds ob- long to lanceolate in outline, green to yellowish green, often flushed with red, 1-2 cm long, 4-5 mm thick; apices of the sepals 1-2 mm long, erect or divergent. Petals very broadlly obovate, 1.5-2.5 cm long. Anthers 6-8 mm long. Filaments 9-12 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 2.54.5 cm long. Stigma lobes 3-5 mm long. Ovary 1.5-2 cm long. Capsule 2.5-4 cm long, 3-5 mm thick. Seeds elliptic in outline, 1.4-2 mm long, 0.6-0.8 mm thick. Self-pollinating complex heterozygote. Gametic chromosome number, n=7 (ring of 14*, ring of 12 and 1 bivalent**, ring of 8 and ring of 6*** or ring of 10 and 2 bivalents**** at meiotic metaphase I). Flowering time: Northern area, November-April; southern area, November-March. Lectotype: Argentina, Prov. Santa Cruz, Rio near Lago Argentino (Lago Santa Cruz), 15 Feb. 1879, E. Ibar (2187) (SGO-041392, GH photograph). Distribution (Fig. 232): Most frequent along the western foothills of the Andes in the provinces of San Juan, Mendoza, Neuquén, Rio Negro, Chubut, and Santa Cruz in Argentina, but with isolated stations in the provinces of Acon- cagua, Santiago, Aisén, and Magellanes in Chile, and very widely scattered lo- calities in the provinces of Cérdoba, San Luis, and Buenos Aires (Bahia Blanca), Argentina. Specimens examined aus 3 plants: ARGENTINA. SAN JUAN: Iglesias, Tocota, 2,480 m, Ruizthal in 1962* (DUSS, MO). MENDOZA: Stony slopes at 9 7 B " m of Punta de Vacas, 2,500 m, Santarius 1459*, 1461, 1464, 1467*, 1475 (DUSS; 1459, 1467 also M; 1467 also MO). Dry rivulet bed near Ruta 7. 4.5 km E of Punta de Vacas, 2 .450 m, Santarius 1476*, 1480, 1483, 1485. 1489*, 1490, 1492*, 1496 (DUSS; 1480, 1489 also CTES, M, MO). Stony slope along Ruta 7, 1 km W of Polva- redas, 2,450 m, Santarius 1504*, 1506, 1513*, 1518 (DUSS; 1504 also M B Arroyo Polvare- das E of Polv aredas, 2,400 m, Santarius 1523*, 1525, 1526*, 1528*, on 1534, 1536, 1540, 1541*, 1544, 1548* (DUSS; 1526, 1528, 1541, 1544, 1548 also CTES, MO). Waste and stony places ca. 2 km N of Tupungato 1,250 m, Santarius 1562* DUE Rocks and stony slopes of the 3 near Villavicencio, 12 km above Villavicencio on Ruta 7, 2,600 m, Santarius 1577*, 1578, 1579*, 1580 ( DUSS; 1579 also CTES, M, MO). 10.5 km above Villa- vicencio, 2 500 m Santarius 1581*, 1582, 1584*, 1588*, 1589*, 1591*, 1592, 1593*, 1595 (DUSS; 1581, 1584, 1588, 1591, 1592, 1593, also CTES, M, MO). 9.5 in above Villavicencio at km 1160, 2,350 m, Santarius 1597*, 1598, 1599*, 1602*, 1603*, 1607, 1608 (DUSS; 1598, 1599, 1602 also CTES, M; 1598, 1599, 1602, 1607 also MO). 9 km above Villavicencio, 2,300 m, Santarius 1610*, 1616, 1621*, 1622*, 1625* (DUSS; 1621, 1622, 1625 also CTES, M, MO). 5 km above Villavicencio (Los Surtidores), 2,050 m, Santarius 1638****, 1639, 1641, 1642, 1643*, 1646*, 1649 (DUSS; 1646 also M, MO). Villavicencio, 1,700 m, Santarius 1652*, 1653, 1654 (DUSS; 1653 also CTES, M, MO). Uspallata, Hecht 1964-79* (DUSS). Nxku- QuÉN: Stony places at Río Limay, 6 km E of Piedra del Águila, Santarius 607*, 609* (DUSS, M, MO; 607 also CTES). Stony places near the ferry across Río Limay at Ruta 237, ca. 75 590 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 km SSW of Piedra del Águila, 150 rp 645* (CTES, DUSS, M, MO, SP). rio NEGRO: San Carlos de Bariloche, Stubbe in 1961* ( CTES, DUSS, M, MO). Sandy and stony waste places at the shore of the Lago 1 Huapi, near railroad station of Bariloche, 780 m, 5 646, 647, 649*, 652, 655, 656", 657, 659, 661, 663*, 668 *, 674, 676, 677, 692, 694, 7 706*, 709, 734, 736, 744, 751 (DUSS; 649, 657, 668, 677, 706 also CTES; 646, 649, 657, ps 677, 706 also M; 646, 649, 657, 668, 676, 677, 706 also MO). N slope of Cerro Otto, ca. 3 km W of Bariloche, along road to tha top on volcanic ashes, 850 m, Santarius 784*, 786, 789*, 796, 797 ( DUSS; 784, 789 also CTES, M, MO). El Bolsón, 325 m, Santarius 838, 840*, 842, 847, 851 (DUSS; 840 also CTES, MO). NW slope at km 17.5 of the road from Bariloche to Villa Catedral, 1.5 km N of Villa Catedral, 900 m, Santarius 853* (DUSS, MO) Meadows about 11.5 km SSW of El Bolsón on the road to Lago Puelo, 300 m, Santarius 821* (DUSS). Airport T Esquel, 675 m, Santarius 903*, 904, 905, 907*, 910, 912 (DUSS; 903, 907 also CTES, M, MO); ibid., Santarius 1393*, 1394, 1398*, 1404 (DUSS; 1393 also M, MO). SANTA CRUZ: Dry Biart bed 2 km N of bridge on Ruta 3 across Rio Gallegos, Santarius 997* (DUSS). Dep. Lago Argentino, Parque Nacional Los Glaciares, N slopes near Guardabosque about 48 km WSW of Calafate, 200 m, Santarius 1067*, 1070, 1074-1076, 1081*, 1086*, 1088, 1089*, ie (DUSS; 1086 also M; 1081, 1086 also MO). Above the campgrounds ca. 51 km WSW of Calafate, 250 m, a lie: 1095*, 1102, 1104, 1108, 1111, 1116, 1121, 1122, 1126 Do 1104 A MO). Road c .lkmS of Punta Bandera near Lago Argentino, 200 m, Santarius 1128*, 1130 (DUSS). f to Punta Bandera near fork to Ventisquero Moreno, 250 m, Santara 1133*, 1135, 1138, 1147*, 1148, 1150, 1159 (DUSS; 1133 also MO). Stony and sandy places near Perito Moreno, 380 m, Santarius 1254*, 1255, 1260, 1268*, 1269, 1270*, ** (DUSS; 1254 also MO). 4 km W of Estancia Las Chilcas, ca, 46 km W of Perito Moreno, 250 m, Santarius 1338*, 1339, 1344, 1345 (DUSS). Río Los Antiguos W of Los Antiguos, 220 m, Santarius 1303*, 1304- 1306, 1309, 1311, 1315*, 1316*, 1317, 1320*, 1321, 1322, 1323* (DUSS; 1306 also CTES; 1303, 1306, 1316 alse M; 1303, 1306, 1320, 1322 also MO). C AISEN: Puerto Ibanez at Lago Buenos Aires, on stony places on the sea shore and slopes up to 300 m above lake, Nodt in 1961 (DUSS). Waste places, shore of Lago Buenos Aires in Chile Chico, 210 m, Santarius 1286*, 1287, 1290, 1292, 1295****, pod Pa ( DUSS; 1286 also M, MO). MAGELLANEs: Road 0-4 km E of Puente Lago Amarga, W of Lago Sar- miento and S of Lago Nordenskjold, 150 m, Santarius 998*, 1001*, 1004*, 1007, 1011, 1014, 1023*, 1024, onse. 1028, 1036, 1037, 1041 (DUSS; 998, 1001, 1023, 1025 also M; 998, 1001, 1004 also M Represe 9 — examined: ARGENTINA. SAN JUAN: Bajada de A Vieja, Hossens 2578 (CORD). Valley of Rio de Los Chupadores, 1 2537 (CORD). Leoncito, Schegaray in 1876 (CORD). Dep. Sarmiento, El Federal, Cuezzo 1705 (LIL, RSA). Arroyo los Dos Puentes sis etm Spe- zzini 231 (BAB). Road from oe to Agua Negra, 3,100 m, Fabris et al. 8383 (L MENDOZA: Villavicencio, 2,900 1 Boelcke 9955 ( BAA, BAB); uin: 19424 (GH, MICH, UC, US); O’Donell 1019 (LIL). Roig 5305 (CORD); Sparre 1498 (S); Nicora 4334 (SI). Puente del Inca, Kurtz 3504 (CORD). Punta de Vacas, Haumann in 1918 (BA). Las Heras, riders del Toro, Lourteig 816 (LIL, NY). Dep. Luján, Est. El Salto, 2,950 m, Ruiz Leal 6183 ' (Leal). bs Malallue, Los Molles, 1,850-1,950 m, e: Leal Lf (Leal) Tupun- gato, Ruiz Leal 2784a (L OM). San Rafael, Minacar at Río Grande, m, Lourteig 734 (LIL). 2 5 . Gillies (K). Cerro Nerado. Ruiz Leal pe (Leal, POM ). Arroyo Mangas in valley of Río Atuel, 1,900 m, Witze 411 (G, US, Z). Banks of Río Atuel S near El Sosneado, Burkart et al. in 1942 I). Re fugio Gral, Alvarado near San Carlos, Cuezzo & Barkley 20Mz475 (LIL). Campo de los Andes near Tunuyan, 1,800 m, Araque 1126 (LIL) sral. Gutierrez near Maipú, Ruiz Leal 25-210 OM). NEUQUÉN ul, & Jo 397 (LP, NY). Est. La Primavera, Castellanos in 1938 (RSA). Between Pulmari and China Muerta, Maldonado 669 (F). Between Norquin and Codihue, Kurtz 6287 (CORD). Las Lajas, Spegazzini 100 (BAB). Lago Nonthué, between Puerto de Gendarmeria and Arroyo near Hua Hum, Valla 3286 (BAA). Paso Flores at Río Limay, in 1938 (LIL-80006). Alu- miné, Giacobbi 12935 (BAA, POM). Santa Maria at Lago Nahuel Huapi, Liunguer in 1934 ). RÍO NEGRO: San Carlos de Bariloche, Lesse 19 (P); Buchtien 1356 (BREM pro parte, GH pro parte); Meyer 7543 (LIL, NY). Cerro Otto, De Barba 945 (LIL, RSA); Boelcke 1702 (CTES). Arroyo Nireco near Bariloche, Meyer 8060 (LIL). Laguna Cari Lauquén, Kurtz 6110 (CORD). El Bolsón, De Barba 422 (F, LIL). Perito Moreno, Novatti 10 (LP). Gral. Conesa, Meyer 7119 (LIL). Choele Choel, 152 m, O'Donell 795 (LIL). CHUBUT: 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 591 Between El Bolsón and Colonia 16 de Febrero, Illin 237 (BR, CORD, HBG, SI). Tehuelches, between Gob. Costa and José de San Martín, Moreau 3619 (BAB). Valley of Rio Chubut, Koslowsky 31-1643 (POM). Est. Leleque near Cushamen, Meyer 7776 (GH, LIL). cey, Kühnemann 648 (RSA). Cordillera, Rio Corcovado, Illin 40 (UC). Dep. Escalante, Est. Los Manantiales near Cañadón Pilar, 380 m, Eyerdam et al. 23800 (G, GH, MO, S, UC). Lago Epuyén, Soriano 1365 (BAA). Esquel, Garcés 286 (SI). SANTA cruz: Rio Gallegos, Tauber 89 (BR). Canadén León, Cittadini 20 (BAB). Puerto Deseado, Correa 3300 (BAA, AB, MO). Güer Aike, O'Donell 4070 (LIL, RSA). Lago Viedma, 300 m, Donat in 1932 (POM). Río Oro, Donat 335 (SI). Lago Posadas, 200 m, Donat 256 (BM, F, G, GH, HBG, K, LIL, MO, NY, S, SI, UC, Z). Junction of Rio Blanco and Rio D = 720 m, Luti 3726 (CORD). Calafate at Lago Argentino, 5 dy e 607 (SI). Between Lago Argen- tino and Lago Viedma, Eyerdam et al. 24347 (G K, UC). Lago 5 esa Aires, Skottsberg ole coRDOBA: El Durazno near 11 y hee & Sleumer 15669 (LIL). Pampa de Achal rd 97 15533 (LIL). Copina, Burkart 7326 (LIL). Valle de los aede 5 ps n 1920 (LIL). Quebrada Ser ae near San Javier, Bridarolli 1582 (LP). Luis: Sierra de E El Rin Hunziker 11828 (CORD). El Morro, 1, 100 n Hunziker 12596 ( CORD); — in 1913 (SI-4671). El Rincón, Conrad & Dietri ich 144, 146, 148, 150, 153 (DUSS). suenos Aires: Bahia Blanca, Claraz 218 (G). CHILE. ACONCAGUA: Intl 2,200 m, Buchtien in 1903 (BM, BREM, GH, L, LY, M, SI, US, W). Road to Argentina, at Rio Blanco, Nicora 4397 (SI). SANTIAGO: Rio Vaso near Romeral, Biese in 1944 (LIL). San Gabriel, 21500 m, Montes 527 (K, MO). AlsEN: Valle Coihaiaue, Rentzell 6129 pro parte (GH). El Paine, Paschke 12246 (CONC). Salto Grande del Paine, a: 2342 (CONC). 1 at Lago Buenos Aires, Heim in 1939 (Z) Chile Chico at L. B. A., Pfister 18480 (CONC). Valle Ibanez, Belem 22799 (CONC). Aisén, Dusén in 1897 icone 486 (S). MAGELLANES: Lago Sarmiento near Puerto Natales, Riie in 1958 (P). Ultima Esperanza, Magens in 1954 (CONC). Laguna Mantecón near Funta Arenas, Cekalovic in 1950 (CONC). Isla 5 I, Gusinde 189 (W). Botanical Garden Kew, England, in 1873 (K; as O. biennis var. undulata). This species derives its genomes from O. santarii and O. odorata. Experi- mental hybridization with these species has established the origin of O. magel- lanica beyond any doubt. Together with O. odorata, O. magellanica is the most frequent species of this section in southern South America, and the two often grow together. It can be distinguished from O. odorata by an array of the char- acteristics it has obtained from O. santarii: upright habit, heavier stems, smaller ‘lowers. From O. santarii it can easily be distinguished by its longer fruits. On the basis of a general similarity in habit, O. magellanica has often been confused with O. stricta, but of course that complex heterozygote has derived both of its genomes within series Allochroa. The short bracts of O. stricta, ulti- mately derived from O. ravenii, immediately separate it from O. magellanica. The pattern of variation in O. magellanica suggests that backcrossing with O. santarii and O. odorata, with the consequent introduction of genetic material into O. magellanica, has been fairly frequent. Within O. magellanica, however, plants with anything except a complete ring of 14 chromosomes are very rare, suggesting a very high level of selection for the restoration of complex heterozy- gosity following hybridization. Where the range of O. magellanica overlaps that of O. santarii, the variation pattern of the former converges on that of the latter, and it begins to resemble series Renneria in general. The farther south one goes, however, the stronger the influence of O. odorata on the populations of O. ma- gellanica. This influence is expressed strongly in the capsules, which become longer and more similar to those of O. odorata southward. The process does not go so far, however, that the boundary between the two is blurred; O. magella- GARDEN [Vor. 64 ANNALS OF THE MISSOURI BOTANICAL o 100 200 300 400 $00 BOO ML ES o 200 400 eoo 800 KILOMETERS so FicunE 240. Range of Oenothera siambonensis (dots). 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 593 nica can always be separated easily from O. odorata because of its more com- pact inflorescence and sturdier habit. Some herbarium specimens are difficult to determine if they include insufficient or immature material; but in living plants, especially under uniform conditions of cultivation, there is never any oubt 36. Oenothera villaricae Dietrich, sp. nov.—Fics. 4, 88, 157, 205. . berteriana “Erlangen” Haustein, Z. Indukt. Abstammungs- Vererbungsl. 84: 418. 1952; Cleland, Jap. J. Genet. 43: 332. 1968. erba annua vel biennis, erecta, rosulata, simplex vel caulis principalis ramosus et ramis MES vel oblique e rosula ascendentibus, 5-10(-15) dm alta. Plantae densissime vel sparse 0 et 5 vel sparse villosae. Folia rosulae anguste elliptica vel oblanceolata, ac E age 8-15 cm longa, 1-2 cm lata; bractea lanceolata vel anguste ovata, acuta, basi truncata, sessilia, praecipue superiora rubro-marginata apiceque incurvata, qu aa ‘subtena brevi- ora vel ad eas subaequalia, raro longiora, 2-4(—6) cm longa, 0.8-1.5 cm lata; folia plerumque nibus exigue ane irregulariter obtuseque serrata. Ioforasoeni plerumque simplex. Tubus floralis 2-3 cm longus. emmae ambito oblongae vel lanceolatae, virides vel flavo- virentes, saepe rubrae, plerumque junctura sepalorum tubo florali anguste rubro-fasciatae; api- ces sepalorum 2-3 mm longi, erecti vel divergentes. Petala latissime obovata, interdum basi pallide rubro-maculata, 1.5-2 cm longa. Stylus brevis, stigmate sub anthesi antheris circum- ato. Ovarium 1.3-2 cm longum. Capsula 2-3 cm longa, 3—4 mm crassa. Semina ambito late elliptica, 1. 1 L 5 mm longa, 0.5-0.7 mm crassa. Numerus gameticus chromosomaticus, n = 7; planta chromosomatice heterozygotica complexa. Erect annual or biennial herb, forming a rosette, unbranched or with a branched main stem and arcuate or obliquely ascending side branches from the rosette, 5-10(-15) dm tall. Plants very thickly to sparsely strigillose and mod- erately to sparsely villous. Rosette leaves narrowly elliptic to oblanceolate, acute, gradually narrowed to the short petiole or subsessile, 10-20 cm long, 1-2 cm wide; cauline leaves narrowly elliptic to narrowly lanceolate, acute, acute to truncate at the base, sessile, 8-15 cm long, 1-2 cm wide; bracts lanceolate to narrowly ovate, acute, truncate at the base, sessile, especially the upper ones with red margins and an incurved tip, shorter than the capsules they subtend or subequal to them, rarely longer, 2—4(-6) cm long, 0.8-1.5 cm wide; leaves mostly weakly undulate at the margins, irregularly serrate with blunt teeth. Inflores- cence usually unbranched. Floral tube 2-3 cm long. Buds oblong to lanceolate in outline, 1.2-1.8 cm long, 4-6 mm wide, green or yellowish green, often flushed with red, usually red striped at the junction of the sepals with the floral tube; apices of the sepals 2-3 mm long, erect or divergent. Petals very broadly ob- ovate, sometimes with a weak basal red spot on each one, 1.5-2 cm long. An- thers 6-7 mm long. Filaments 7-11 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 3—4.5 cm long. Stigma lobes 3-4 mm long. Ovary 1.3-2 cm long. Capsule 2-3 cm long, 3-4 mm thick. Seeds broadly elliptic in outline, 1.1-1.5 mm long, 0.5-0.7 mm thick. Self-pollinating complex heterozygote. Gametic chromosome number, n = 7 (ring of 14* at meiotic meta- phase I). Flowering time: November-March. 594 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Type: Grown from seeds and cultivated in the Botanical Garden of Düssel- dorf, Germany, 2 Aug. 1972. Source: Chile, Prov. Cautín, Molco at Lago Villa- rica, end of 1960, W. Stubbe (MO-2155226, holotype; CTES, DUSS, M, isotypes). Distribution (Fig. 234): In the Chilean provinces of Cautín, Valdívia, Osorno, Llanquihue, and on Isla Chiloe, as well as in a few localities on the eastern slope of the Andes in the Argentine provinces of Neuquén, Río Negro, and Chubut. Specimens examined from es plants: CHILE. CAUTIN: Sitio Göpfert at Lago Villarica, meadow above the seashore between Antumalal and Pucón, Gépel in 1961* (CTES, DUSS, M, MO). Laguna Verde at Río Allipén near Volcán Llaima, Gépel in 1961* (CTES, DUSS, M, MO). Molco at Lago Villarica, Stubbe in 1960* ( DUSS, M, MO). VALDIVIA: Taco Tres at Río San Pedro near Lago Rinihue, Stubbe in 1961* (CTES, DUSS, M, MO). Las Animas near Valdivia, Göpel in 1961* ( CTES, DUSS, M, MO). Isla Teja in Valdivia, Stubbe in 1960* (CTES, DUSS, M, MO). Collico near Val- divia, Koch in 1960* (DUSS). Along the railroad near Antilhue, Stubbe in 1961* (CTES, DUSS, M, MO). osorno: Stony places at S edge of Osorno, Stubbe in 1961* (CTES, DUSS, M MO). Pan-Amer. Highway, 5 km S Valdivia-Osorno boundary, Wiens in 1967* (CTES, S, MO). ARGENTINA. NEUQUÉN: Pucara in the Parque Nacional Lanín, Schachovsky in 1965* (DUSS). cnonur: Meadows ca. 11.5 km SSW of El Bolsón along road to Lago ess 300 m, Santarius 823*, 824, 825*, 831*, 833 (DUSS; 823 also CTES, M; 823, 833 also M TIVATED: O. "berteriana" from the Botanical Garden of Erlangen in cue re- ceived in 1960* (CTES, DUSS, Ms MO). Additional specimens exam ned: CHILE. CAUTIN: Calvert in | 1914 (BM). Freire, Claude-Joseph 5899 (US). Río Zuapa, enu in 1905 (G). VALDIVIA: Valdivia, Lechler (M); Hollermeyer 56 (S); Buchtien m (US), in 1898 (US); Gay 81 (P). Railroad to e asi n in 1904 (M Panguipulli Gay in 1834 (G, SGO). osorno: Lago Llanquihue, Calvert in 1912 (BM). ' CHI- nal de Dalcahue, Funck 126 (P). Without exact locality: 1 91 (HBG). 5 NEUQUÉN: Lago Quillén, Dawson & Schwabe 2874 (BAB). mío NEGRO: Bariloche, Fabris 1124 (M). El Bolsón, Meyer 7884 pro parte (NY); Illin 6875 (BAB). Specimen from plants cultivated in gar Botanical Garden of Valence, Italy, 9 eli of the Botanical Garden at 5770 under the name O. berteriana Spach, plant of Chile, 1923, Herb. E. J. 5 14-140 (MPU). This specimen is identical with the strain known to geneticists as O. berteriana. oe specimen was seen from the Botanical Garden Bremen, Germany, Fahrenholtz jn. 1923 (BREM). This newly proposed species has been known to geneticists for some time, since it is identical with the “Erlangen” strain said to be O. berteriana. The original provenance of this line is unknown, but presumably it came from one of the provinces of Chile named above. Oenothera berteriana itself is a synonym of O. affinis. Like O. magellanica, most herbarium specimens of O. villaricae are identi- fied as O. stricta, which of course had a different origin. The Renneria genome of O. villaricae was once more contributed by O. santarii, whereas its Allochroa genome is derived from O. ravenii (see also Cleland, 1968). Since, as has been pointed out several times, Chile has no native chromosomally homozygous spe- cies of the subsection (except for O. coquimbensis), and since the main area of O. villaricae seems rather clearly to be Chilean, it probably originated following hybridization between O. magellanica and O. stricta. ybrids between O. santarii and O. villaricae consist of two classes, one like each parent. Hybridization with O. ravenii is only possible if that spe- cies is used as the male parent; the plastids of chromosomally homozygous O. ravenii from Brazil do not function in a genetic background of O. villaricae. DIETRICH—SOUTH AMERICAN OENOTHERA 1977] 595 soo oo MILES eoo KILOMETERS 40 mE 30 [T] 80 J 60 i so Fıcure 241. somal complex in O. magellanica and O. villaricae (circle Range of homozygous Oenothera santarii (filled triangles) and as a chromo- s). 596 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 They are colorless, and hybrids between these two species are viable only when there are sufficient green plastids derived from the O. villaricae parent. This is usually only the case when O. villaricae is the female in the hybridizations. One of the classes of F; hybrids between O. villaricae and O. ravenii is iden- tical to O. ravenii, but the second class does not exactly match O. villaricae. This is probably because the large flower size of O. ravenii is dominant in O. villaricae, and also because the O. ravenii complex in O. villaricae is derived via O. stricta and probably considerably altered in the course of evolution of that species. Oenothera villaricae can be distinguished from O. magellanica by its shorter, red-margined bracts and shorter capsules. Both species differ from O. stricta in their upright, sturdy habit and more compact inflorescence. The Argentinan plants of O. villaricae are somewhat different from the Chil- ean ones in that their bracts are usually shorter. This would seem to indicate a stronger influence of the O. ravenii genome. In addition, they have a somewhat more delicate habit and are consistently taller when grown under uniform cul- tural conditions. It does not seem desirable to separate them taxonomically, however, because they have had a common origin and are closely related. If all of the small differences within the section that are preserved by the syndrome of self-pollination and complex heterozygosity, and which have in many cases come to characterize populations, were to be recognized taxonomically, one could recognize literally hundreds of species without any gain whatsoever in taxonomic utility. Taxonomy serves its end better as a device for summarizing and grouping information about a particular group of organisms, not by splitting them into an excessive number of categories. 37. Oenothera hechtii Dietrich, sp. nov.—Fics. 89-91, 158. O. parodiana "Vila Nougues" Hecht, Indiana Univ. Publ. Sci. Ser. 16: 277. 1950; J. Hered. 99. 1970. 61: erba annua vel biennis, erecta, rosulata, simplex vel caulis principalis parce ramosus et ramis us arcuato e rosula ascen entibus —10 dm alta. Plantae parce vel sparse strigulosae c n : elliptica, acuta, basi acuta vel 5 sessilia, 6-12 cm longa, 1-2 cm lata; bractea a irregulariter serrulata. (ea sim jux. Tubus floralis 4-5.5 cm n longus; cae Acre lanceolatae, flavo-virentes, junctura sepalorum tubo florali rubro-fasciatae, 1.5-1.8 cm longae, 4-6 mm crassae; apices sepalorum ca. 1.5 mm longi, erecti. Petala obovata, 1.8-2.2 cm longa. Stylus brevis, stigmate sub anthesi antheris circumdato. Ovarium 1-1.3 cm longum. Capsula 2.5-3 cm longa, ca. 3 mm crassa, extremi valvarum distincti, discreti, crenatique. Semi 1- = late elliptica, 0.9-1.1 mm longa, 0.6-0.7 mm crassa. Numerus gameticus mao. = 7; planta chromosomatice heterozygotica complexa Erect annual or biennial herb, anes a rosette, with simple or little- branched main stem and widely arching side branches arising from the rosette, 5-10 dm tall. Plant moderately to sparsely strigillose and villous, more densely so in the region of the inflorescence. Rosette leaves very narrowly elliptic, acute, attenuate at the base, sessile, 12-16 cm long, 2-2.5 cm wide; cauline leaves nar- rowly oblong to narrowly elliptic, acute, acute to rounded at the base, ses- 1977] DIETRICH—SOUTH AMERICAN OENOTHERA 597 sile, 6-12 cm long, 1-2 cm wide; bracts lanceolate, acute, rounded to truncate at the base, sessile, 2.5-3 cm long, 0.8-1.2 cm wide; leaves plane at the mar- gins, irregularly serrulate. Inflorescence unbranched. Floral tube 4-5.5 cm long. Buds lanceolate in outline, yellowish green, red striped at the junction of the sepals with the floral tube, 1.5-1.8 cm long, 4-6 cm thick; apices of the sepals ca. 1.5 mm long, erect. Petals obovate to broadly obovate, 1.8-2.2 cm long. An- thers 5-6 mm long. Filaments 10-13 mm long. Style short, the anthers shedding pollen directly on the stigma at anthesis, 5-7 cm long. Stigma lobes 4-5 mm long. Ovary 1-1.3 cm long. Capsule 2.5-3 cm long, ca. 3 mm thick, with distinct, free, crenate ends to the valves. Seeds broadly elliptic in outline, 0.9-1.1 mm long, 0.6-0.7 mm thick. Self-pollinating complex heterozygote. Gametic chro- mosome number, n = 7 (ring of 14* at meiotic metaphase I Type: Grown from seeds and cultivated in the Botawical Garden of Diissel- dorf, Germany, 4 Aug. 1972. Source: Argentina, Prov. Tucuman, Villa Nougés near Tucumán, A. Hecht 1964-81 (MO-2155202, holotype; CTES, DUSS, M, isotypes). Distribution (Fig. 235): So far known only from the type locality. Specimen examined from cultivated plan ARGENTINA. Prov. Tucumán, Villa Rand near Tucuman, Hecht 1964-81* (DUSS, M, MO). This new species is dedicated to Adolph Hecht (1914-), student of Oeno- thera. Because of its superficial similarity to O. parodiana subsp. brasiliensis, it was not realized at first that O. hechtii belonged to series Clelandia. Conse- quently it has not yet been possible to test its parentage by experimental hy- bridization, although some deductions can be made on a morphological basis. Clearly, the Allochroa element in O. hechtii can have been derived only from O. ravenii. The rosettes of O. hechtii and its flowers, which have relatively long tubes and sometimes overtop the stem, are reminiscent of those of O. longituba of series Renneria 38. Oenothera elongata Rusby, Mem. Torrey Bot. Club 3(3): 33. 1893.—F'cs. 159, O. serratifolia Krause, Repert. Spec. Nov. Regni Veg. 1: 168. 1905. type: Bolivia, Dep. Tarija, Toldos near Bermejo, in a canyon, ca. 2,000 m, 9 Dec. 1903, K. Fiebrig 2374 (B, de- stroyed in World War II, UC 292 and 689 fragments and vhotograph, F, GH and POM photographs; G, isotype). Erect annual herb, not forming a rosette, unbranched or branched from the base upward, 6-15 dm tall. Plants densely to sparsely strigillose, densely to sparsely long- and short-villous, and densely to sparsely glandular-pubescent. Cauline leaves very narrowly elliptic to lanceolate, acute, attenuate to rounded at the base, 5-12 cm long, 0.5-1.5 cm wide; bracts lanceolate to narrowly ovate, acute, 2.5-5 cm long, 1-1.5 cm wide, longer than the capsules they subtend; leaves plane at the margins and remotely serrate, the teeth blunt. Inflorescence simple or branched. Floral tube 7-10 cm long. Buds narrowly oblong to lanceo- late in outline, 1.5-2.5 cm long, 4-6 mm thick, often red striped at the junction 598 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 40 80 T l - | | + {10 10 | “° l | | " | ` 6 t ° " ç / S ç M o I / id j ES L Si 3 i , ‘ f x IR ` l : Cx Pu CAD t so dii BEL. A. 10 / \ 1 S as ' h | 20 20 | - | Z | | | | | 30} 30 l | 40 Le 400 s 400 MILES. M T — 0 200 400 eoo 800 KILOMETERS / so 4 7 iso / / / / | / / / f / / oo 4257, 445K, 537F, 614. var. limensis var. nocturna om, 582, 613 r. pubescen inan ane 616 s 446K, 457, 467F, 4887, 4897, 31F, 548F, 6111 longiflora 4341, 4367, 4377, 446K, 4667, 941, 503, 5067, 509, 5141, 5271, 5301, E 5491, 71 5647, 5651, 5681, 5707, 5747, 6011 Es jail 459F, 486F, 5107, 513+, 535F, 546F, 559F E. bs 426+, 459F, 486F, 5107, 511K, 5111, 511, 5411, 546F, F 561 X stricta 514 longituba 434t, 4391, 4407, 4437, 446K, 456F, 463, 469F, 4807 4827, 4851, 531F, 544F, 554F, 5977, 5997 macrosiphon 524 as 4281, 4347, 439+, 449K, 4637, 5007, 512F, 569F, 587, 5947, 2725 5967, 598F var. chubutensis 587 malacophylla Soie pe oer pan 347, 447K, E 479F, 492, 5177, Eod 5361, 546F 5491, 551F, 5651, 5667, 5687, 5701 micans 618 mollissima 4267, 4397, 448K, 460F, 4907, 530, 535F, 5437, 546F, 552, 553, 556, 5587, 5607, 5627, 569F, 587, 602, 607F, 6227 —subsp. odorata 5 9491 —su P Lon —var. grandiflora 524 —var. sabulosa 5407 DIETRICH—SOUTH AMERICAN OENOTHERA 625 r. valdiviana 538 —var. villosa 530 5 hec 4927, 499F, 519f, 60, 578F, 60 nana 428+, 4407, d 4441, 445K, 4511, „ 4617, 477F, 487, 544F, 569F, nervosa 514f ii 447K, 4907, 510F, 537F, 546F, , 5821, 582, 588F, 6147 "m. 426%, 4287, 4347, 4361, 4377, 4381, 439t, 440t, 444t, 447K, 458F, 4637, 552, 5521, 5591, 554F, 5581, 565. 568*, 570%, 580. 582, 587, 5914, 227 — var. glaucescens 494 virescens 494 as ah = 494 =Ë. 94 —f, emulate 494 odoratissima 49 parodiana 503, 506, 5237, 5491, 553, 5561, 568, 970, 6011, 566K, 566, 5741, 586F, 598F subsp. strigulosa 447K, 504F, 5651, 566K, 568, 588F “Concordia” 570 “Siambo 602 Villa Nen 596 vb 4397, 446K, , 044F, 561F pe dune x O. indecora subsp. bona- 087 4537, 456F, pennellii 61: peruana = 439}, 446K, 452F, 453, AGTI Ps 4851, 4897, 531F, 548F, 585 picensis 15 ae 1 E 550, 5531, 5561, 5647, 598F E bonariensis oe 5237, 550K, 584F Es ` cordobensis 449K, "d .499F, 546F, 550K, 553, 5651, 58 cae: picensis 499F, 5501, 551K 552, 5567, 5 propinqua 538 var. sparsiflora 538 prostrat pseudoelongata 448K, 4857, 571F, 5871, 607F porno po 501F, 516F, 562, EE 607 oben sqa punae RM Ps 4407, 4437, 445K, 464F, 626 487, 4887, 4897, 523F, 5237, 5247, 535F, 544F, 5857, 586F, 5871, 609 ravenii 4267, 434+, 4367, 4407, 450K, 4637, 466, 494+, 500, 5067, 5097, 5107, 5157, 5237, 5457, 5471, 5537, 5567, 5647, 5657, 5667, 5687, 5747, 5821, 5917, 5947, 5967, 5977, 6017, 604+ subsp. argentinae 449K, 483F, 5031, 503K, F, 603F —subsp. chilensis 459F, 483F, 5031, 503K, 508, 5431, 551F, 603F subsp. ravenii 431t, 447K, 458F, 479F, 481F, 5037, 503K, 503, 5067, 5087, 5097, 531F, 546F, 548F, 603F recurva 446K, 456F, 4657, 473F, 480, 4857, 5577, 509 rivadaviae 445F, 448K, 461F, 491F, 534, 561 F, 598F * ed 446K, 454, 473F, 476, 482, Pom 5437 sandiana 446K, 475F, 4787, 482, 4891, F, 559F, 612 santarii rage 4347, 436T, 4397, 4401, 446K, E F, 461, 469F, 531F, 544F, ee 5941, 595F "ms 15 4347, 4367, 4377, 4397, 445F, 446K, 4537, 456F, 465+, 4687, 4707, 471F. 473F, 474, 4787, 4807, 4827, 4857, 4897, 531 F, 544F, 563F, 6007, 6047, 6057, 6227 sellowiana 5067 serratifolia 597 siambonensis yid p 465F, 516F, 92F, striata 536, 4: ~ stricta 4281, 4367, 4397, 4407, 494, 506, 5437, 5457, 5537, 5807, 587, 5911, 5947, 5967, 598F, 602, 603F, 6187 subsp. altissima 448K, 461F, 495F, , 544, 546F, 5471, 563F —subsp. argentinae R 461F, 495F, 538K, cd stricta Hu. 4281, 450K, 495F, 5147, 5227, 5387, 538, 538K, “San ibi gen 617, 6187, 6197 stuebe 13 1 6177 tafiensis 446K, 4537, 466, 4721, 4747, 601+ —subsp. parviflora 453+, 453F, 468K, »- - e° ~] — "nj e g Ke; o>) a 446K, 456F, 465, ` AT3F, 478, 4817, 4827, 4857, 554F, 5997 tetraptera 533+ tucumanensis 448K, 465F, 520F, 5237, 581F, 604+, 606 M ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ime) 494, 4987, 5307 valdiviana 538 verrucosa 4347, 4367, 4447, 445F, 447K, , 4907, 4927, 504 F, 5177, 535F, a A 5767, 577, 5797, 5851, 586 6147 versicolor 0 4367, 4397, 446K, 4517, 45 547, 454, 4577, 4617, 4651, ye 8 4827, 4857, 4897, 531 F, T villaricae 4287, 4397, 4407, 449K, 4637, , 512F, 544F, 573F, 593, 595F, 6035 villosa 4257, 4981, 5147, 5247, 5301, 5331, 617 subsp. strigosa hg 617 obs 4651, . 565t Onagra 450, 489, 155 arguta 538 fusca 454 mollissima 590 9 Onothera albicaulis var. tigrina subvar. co- 5 quimbensis 5 argentinae 519, 519 197 . 538 e Tonto 511 var. rna 582 var. se scllow i 513 —race ihe esi 530 var. indecora 517 dcn 594 — var. ibari kasara uni, rg 4287, 489, 585, 6127 erteria x 575 i 51 me a aaa 530 nocturna 582 odorata 494 punae 609 Renneria 4257, 4267, 428+, 4327 Viridiarum cliffortianum 5327 WOOD ANATOMY OF ONAGRACEAE: ADDITIONAL SPECIES AND CONCEPTS' SHERWIN CARLQUIST” ABSTRACT vod anatomy of Epilobium colchicum subsp. ag je eeu excorticata, and Hauya heydean is described qualitatively and quantitatively. For the latter two species, large logs vere available and wood portions from both inside and 1 were analyzed. Although these | e species offer no features new for Onagraceae, each adds features new for its respective genus. By means of numerical indices which are termed vulnerability and ABER respec- tively, values are presented to show the range in ecological d of woods of the three species, as well as of all Onagraceae studied earlier. Onagraceae d. a wide range in these indices and probably form a good model of what use indie in families with a broad ecological range will demonstrate. Wood from inside of logs of Fuchsia excorticata and Hauya heydeana is more xeromorphic than wood from the periphery. In my monograph of wood anatomy of Onagraceae (Carlquist, 1975a), I attempted a sampling of woods based largely on availability of portions of suit- able size. In any family in which herbs predominate, one is faced with arbitrary decisions as to whether some species which form little secondary xylem should be included. However, after the appearance of the monograph, Dr. Peter H Raven placed at my disposal three wood samples with abundant secondary xylem for their respective genera, and otherwise cf more than passing interest. MATERIALS AND METHODS Epilobium colchicum Alboff subsp. colchicum (sect. Chamaenerion) was collected by Dr. Peter H. Raven from a streambed near the Lagodekhi Hotel in eastern Georgia, U.S.S.R., and is unusually woody for an Epilobium. The species of Fuchsia that forms perhaps the largest trees in that genus (and perhaps also in the family) is F. excorticata (J. R. & G. Forst.) L. f. A section from the base of a tree approximately 0.9 m in diameter (outline of section irregular) was supplied by the New Zealand Institute of Forestry. Dr. Dennis E. Breedlove's mesoamerican field work yielded a log, approximately 25 cm in diameter, of Hauya heydeana Donnell Smith. Because all the Hauya wood samples utilized in my earlier study were of H. elegans subsp. cornuta, material of the second species of this interesting genus was especially welcome. All three wood samples were dried. Methods of study were the same as those employed for dried sam- ples in the earlier paper. Dr. Larry DeBuhr's work in preparing sections and macerations and in obtaining data is gratefully acknowledged. Because differ- ences in wood anatomy were observed (Carlquist, 1975a) in samples of differ- ent diameter in Hauya elegans subsp. cornuta, both inner and outer portions of 1 This study has been aided by a grant from the National Science Foundation, BMS 73- 07055-Al. The wood of Fuchsia excorticata was provided by Dr. E. J. Godley, D.S.LR., Christchurch, be w Zealand. 2 Claremont Graduate School, Pomona College, and Rancho Santa Ana Botanic Garden, Claremont, 25 ae 91711 Ann. Missouni Bor. GARD. 64: 627-637. 1977. 628 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 v e a” e 0 e. 4 ata HY : jx vé du — * è CERCEL "Y. ` MAN FiGungs 1—4. Wood sections of Epilobium me Hauya.—1-3. Epilobium colchicum Alboff subsp. colchicum, Raven 26519 Sean —]. Transection; parenchyma bands not shown in this photograph.—2. Tangential section; rays few, inconspicuous.—3. Portion of vessel wall from tangential section.—4. Hauya 1 and Donnell Smith, Breedlove 15653 (MO). Radial section of ray cells showing nature of walls and starch grains embedded in dark-staining amor- phous materials. [Magnification indicated by atl raph of stage r. enlarged a same scale as applic: able photomicrographs. Figs. 1-2, : gar above Fig. 2 (finest divisions = 10 um). Figs. 3—4, scale above Fig. 4 cee = 10 um).] et 1977] CARLQUIST—WOOD ANATOMY OF ONAGRACEAE 629 the logs of Fuchsia excorticata and Hauya heydeana were studied. Quantitative data for both portions are reported below. Qualitative features of inner and outer portions are the same unless otherwise mentioned. ANATOMICAL DESCRIPTIONS Epilobium colchicum subsp. colchicum, Raven 26519 (MO), Figs. 1-3. Growth rings present as parenchyma bands that are discontinuous in places around the stem. Mean vessel diameter, 50 um; mean vessel element length, 184 um. Vessels mostly solitary (Fig. 1); if grouped, in radial chains or multiples; mean number of vessels per group 1.36. Mean number of vessels per mm? of transection 38.1. Perforation plates simple. Lateral wall pitting of vessels basi- cally alternate, appearing somewhat scalariform because pits are laterally elon- gate (Fig. 3). Pits conspicuously vestured (Fig. 3). Mean libriform fiber length 277 um. Mean libriform fiber wall thickness 1.6 um. Libriform fiber walls not gelatinous; pits simple. Axial parenchyma in the form of bands, with also a few vasicentric cells forming strands of one to three cells. Bands of axial parenchyma probably contain interxylary phloem, but determination uncertain because of lack of liquid preservation. Multiseriate rays more frequent than uniseriates, but both inconspicuous (Fig. 2) because upright cells predominate in multiseriates, with only a few square and procumbent cells. Uniseriate rays consist of upright cells only. Mean height of multiseriate rays 567 jum; uniseriates, 84 wm. Ray cells thin to moderately thick, the latter sometimes with bordered pits. No crystals observed. Amorphous deposits of dark-staining materials in some ray cells ( Fig. 1). Wood not storied. Fuchsia excorticata, Figs. 5-8. Growth rings inconspicuous in inner wood (Fig. 5), with narrower vessels in latewood, wider vessels in earlywood. Growt rings not evident in outer wood. Mean vessel diameter 73 wm in outer wood, 63 um in inner wood. Mean vessel element length 325 um in outer wood, 259 Im in inner wood. Vessels solitary or in multiples (Fig. 5), averaging 1.52 per group in outer wood, 1.76 per group in inner wood. Mean number of vessels per mm? of transection 32 in outer wood, 57 in inner wood. Perforation plates sim- ple. Tyloses present in vessels, numerous, thin-walled (Fig. 6, right). Lateral wall pitting of vessels (Fig. 8) consists of alternate pits, angular and rhomboidal in outline or laterally elongate. Pits conspicuously vestured. Mean libriform fiber length 598 um in outer wood, 562 um in inner wood. Libriform fiber wal thickness 2.5 um in outer wood, 2.9 um in inner wood. Libriform fiber walls not noticeably gelatinous. Libriform fibers prominently septate (Fig. 7). Ex- tremely minute vestiges of borders observed on pits of some libriform fibers (Fig. 7). Interxylary phloem absent. Axial parenchyma scanty vasicentric; strands consisting of three to five cells per strand. Rays predominantly multi- seriate (Fig. 6); uniseriates infrequent, virtually absent in inner wood. Mean multiseriate ray height 755 jum in outer wood, 494 um in inner wood; uniseriates (outer wood), 224 um. Multiseriates composed of upright, square and procum- bent cells. Uniseriates composed wholly of erect cells. No crystals observed. A few ray cells with dark-staining contents (Fig. 6). Wood not storied. 630 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 2 “...... — "o o 2 — ` = ` sya x x 9.572. š Zz * ons — m $ 4 - - š f - 1 Ld 7 MIO 8 7 " 121775 ; 1> ---- qe t IP EAURS í x P- 123,226 N ;UREs 5-8. Wood Pu of Fuchsia excorticata (J. R. & G. Forst.) L. f., collected by the New Zealand Forestry Institute.—5. Transection, from near center of large log.—6. Tan- gential section, from near S of large log.—7. Septate fibers, showing pitting, from radial section.—8. Vessel wall from radial section, showing vestured pits, dom in dicen some late: rally elongate. [Magnification for Figs. 5-6, shown above Fig. 2. Scale for Fig 7-8, shown above Fig. 4.] 1977] CARLQUIST—WOOD ANATOMY OF ONAGRACEAE 631 Hauya heydeana, Breedlove 15653 (MO), Figs. 4, 9-12. Growth rings incon- spicuous, vessels wider in earlywood (Fig. 9). Mean vessel diameter 111 um in outer wood, 80 um in inner wood. Mean vessel element length 506 um in outer wood, 481 um in inner wood. Vessels mostly grouped in short radial chains (Fig. 9), 2.52 per group in outer wood, 2.24 in inner wood. Mean number of vessels per mm? of transection 31 in outer wood, 56 in inner wood. Perforation plates simple. Lateral wall pitting of vessels alternate, pits round in outline with some tendency toward laterally elongate pits. Vesturing on pits not readily visible. Mean length of libriform fibers 874 um in outer wood, 903 jum in inner wood. Mean libriform fiber wall thickness 2.5 jum in outer wood, 2.0 um in inner wood. Libriform fibers with inner portion of wall markedly gelatinous (Fig. 10). Axial parenchyma mostly vasicentric, but with some cells in tangential bands (Figs. 9, 10, 12), as in H. elegans subsp. cornuta; parenchyma cells consist of two or three cells per strand. No interxylary phloem present. Both multiseriate and uniseriate rays present (Figs. 11-12). Mean multiseriate ray height 497 um in outer wood, 609 um in inner wood. Mean uniseriate ray height 352 um in outer wood, 305 um in inner wood. Both multiseriate and uniseriate rays consisting of upright, square and procumbent cells. Ray cell walls thick (Fig. 4), some with bordered pits. Massive deposits of dark-staining materials in ray cells and axial parenchyma (Figs. 4, 9-10, 12). Starch grains also present in ray cells (Fig. 4) and axial parenchyma cells. Wood not storied. COMPARISONS Epilobium colchicum (sect. Chamaenerion), a woody perennial, differs from E. paniculatum (sect. Xerolobium), a rank annual, most notably in the presence of axial parenchyma bands ( Carlquist, 1975a). Phloem is presumptively present in these bands. Because these bands do not extend all the way around the stem, the apparent absence of cork cells, such as reported by Moss (1936) in Epilobium angustifolium (also of sect. Chamaenerion), is understandable. Epilobium col- chicum shows laterally widened pits on the vessel walls (Fig. 3) with exceptional clarity. These pits are very clearly vestured. Species of the tribe Epilobieae do not, in wideness of vessels, length of vessel elements, and low number of vessels per mm? of transection, approach the two species of Hauya. Still, one may note that E. colchicum has the most mesomorphic woods of the three species of Epi- lobieae now studied. In view of its riparian habitat, which contrasts strongly with the xeric habitats of the other two species, this correlation is logical. Fuchsia excorticata falls within the range of wood features reported for Fuchsia earlier. Scanning electron micrographs of the vestured pits of F. excor- ticata have been presented by Butterfield & Meylan (1973). The potential inter- est of F. excorticata in relation to wood anatomy of Onagraceae is whether, with trunks of such large size, changes in quantitative characteristics occur with age. As the above descriptions indicate, some changes do occur. Vessel elements are appreciably wider, longer, and few per mm”? in the outer wood than in the inner wood. Libriform fibers are longer and slightly more thin-walled in the outer wood. Multiseriate rays are taller in the outer wood. Although ray histology ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Ii * t 9 2 — sd — | a Pi Ld “=. me E ^ du. 00, € zs 9-12. be sections from periphery of log of oum heydeana Donnell Smith i 15653 (MO).—9. Transection, showing a growth rin Transection, enlarged, showing gelatinous in of libriform fiber rs, dark-staining content of 5 cells, distribution of axial parenchyma.—11. Tangential section, showing nature of rays.—12. An- other tangential section, ur massive deposits of roe materials in rays, ixi paren- chyma. [Scale for Figs. 9, 11-12, shown above Fig. 2 ale for Fig. 10 Asi Fig. 10.] 1977] CARLQUIST—WOOD ANATOMY OF ONAGRACEAE 633 does not seem to change, there is a change in presence of uniseriate rays. In the outer wood of F. excorticata (Fig. 6), uniseriates are present, whereas they can be said to be virtually absent in the inner wood. This is exceptional in Fuchsia, although in most species studied, multiseriates do predominate over uniseriates (Carlquist, 1975a). The quantitative changes with age are in accordance with those one expects in a typical woody dicotyledon (Bailey & Tupper, 1918; Carl- quist, 1975b ) Hauya heydeana differs in some features of wood anatomy from the species studied previously, H. elegans subsp. cornuta. Most notable are the ergastic materials. Whereas H. elegans subsp. cornuta had, in all collections, large crys- tals within fibriform cells (sometimes with smaller crystals), no crystals of any sort could be detected in any of the sections of H. heydeana. On the other hand, wood of H. heydeana is dark in color, a fact undoubtedly related to the abun- dance of dark-staining deposits in ray and axial parenchyma cells, as revealed by the photomicrographs (Figs. 4, 9-10, 12). The presence of thick-walled ray cells with bordered pits (Fig. 4) is to be expected in view of the occurrence of this phenomenon in H. elegans subsp. cornuta, Camissonia crassifolia, and Epi- lobium colchicum subsp. colchicum. The occurrence of axial parenchyma cell groups in H. heydeana (Fig. 10) is much like that in H. elegans subsp. cornuta, and emphasizes the generic distinctness of Hauya. Ray histology does not ap- pear to change appreciably with age in H. heydeana, although such large trunks as those of H. elegans subsp. cornuta were not available, perhaps because this species does not form as large trees. The presence of starch in ray cells of II. heydeana (Fig. 4), conspicuously preserved because of embedding of the starch grains in the amorphous deposits, is to be expected for Onagraceae as a whole and was also observed in H. elegans subsp. cornuta. The two species are alike in the absence of septa in fibers, absence of interxylary phloem, and absence of tyloses. The gelatinous fibers of H. heydeana (Fig. 10) differ from the non- gelatinous ones in H. elegans subsp. cornuta, although gelatinous fibers are very common in Onagraceae (Carlquist, 1975a). Pits on vessels of H. heydeana are probably vestured, but that vesturing is so fine that it appears only as a slight darkening of the edge of the pit aperture; scanning electron microscopy would be required for definitive demonstration. Qualitatively, wood characteristics of H. heydeana are much like those of H. elegans subsp. cornuta, indicating an equally mesomorphic conducting sys- tem conformation. The changes from inside to outside of a log of H. heydeana in quantitative features are much like those of H. elegans subsp. cornuta or Fuchsia excorticata: vessels elements are wider, longer, and fewer per mm? of transection in the outer wood. Rays are shorter in the outer wood of both species of Hauya, contrary to the increase in ray height in Fuchsia excorticata. ECOLOGICAL CONCEPTS In my survey of Onagraceae (Carlquist, 1975a), I used, for groupings of species, an index based upon the sum of mean vessel diameter and mean vessel element length. That index appeared to reflect accurately the ecology of onagra- 634 ANNALS OF THE MISSOURI BOTANICAL GARDEN TaAnLE I. Ecological indices for woods of Onagraceae. [Vor. 64 Species V M Fuchsieae Fuchsia boliviana Britton 1.54 559 F. cyrtandroides J. W. Moo 5.15 2,260 F. excorti cata (J. R. & G. Forst.) L. f. 1.08 280 sac 2.28 741 F. un 1.67 499 F. dicic mis Lam. var. globosa (Lindl.) Bailey 1.75 673 F. magellanica var. macrostemma (Ruiz & Pavón) Munz 1.3 489 F. paniculata Lindl. 3.91 2,151 F. parviflora Lindl 0.52 245 F. n Zucc. 1.28 591 F. tincta I. M. Johnston 1.08 356 F. Aion Krause 0.22 71 All Fuchsieae 1.99 634 Lopezieae Lopezia a Zucc. 1.17 579 L. langmaniae Miranda 2.07 797 L. bus yi 9 A Plitmann, Raven & Breedlove 5.79 1,847 L. lopezioides (H. & A.) Plitmann, Raven & Breedlove, Breedlove 7268 2.00 902 E. lopezioides, Breedlove 8052 2.96 929 L. miniata Lag. ex DC. subsp. miniata 0.63 187 L. miniata subsp. paniculata (Seem.) Plitmann, Raven & Breedlove 1.21 439 L. racemosa Cav. subsp. moelchenensis Plitmann, Raven & Breedlove 0.32 86 L. racemosa subsp. racemosa 0.86 237 L. riesenbachia Plitmann, Raven & Breedlove 0.46 155 L. semeiandra Plitmann, Raven & Breedlove 3.68 1,417 Lopezieae: shrubs combined 2.70 961 Lopezieae: annuals, suffrutescent perennials combined 0.67 215 Onagreae Calylophus hartwegii (Benth.) Raven subsp. pubescens (A. Gray) »wner & Raven 0.14 26 C. serrulatus ( Nutt.) Raven 0.13 18 Camissonia californica Raven 0.76 234 C. crassifolia ( Greene) Raven 0.12 25 C. cheiranthifolia ( Hornem. ex Spreng.) Raimann 0.41 68 C. megalantha (Munz) Raven 0.83 338 Clarkia xantiana A. Gray 0.35 119 Gaura biennis L. 0.66 174 G. longiflora Spach 1.10 375 G. parviflora Papa 1.13 267 G. sinuata Nutt. ex Sér. 0.14 15 G. villosa Torr. su d. villos 0.52 134 „ fruticulosus (Benth ) Brandegee subsp. glaber Thomas) Carlquist & Rave 0.53 120 G. rubricaulis Schlecht : Cham. 0.75 271 Heterogaura heterandra bee 0.69 197 pie tle deltoides Tor & Frém. cubes howellii (Munz) Klein 0.81 237 O. V eni te Hooker 0.74 169 O. elat 1.19 353 O. lailia 0.20 48 Xylonagra Abos (Kell.) Donnell Smith & Rose subsp. wigginsii Munz 1.68 314 Onagreae: annuals combined 0.54 161 Onagreae: caudex perennials 0.22 48 1977] CARLQUIST—WOOD ANATOMY OF ONAGRACEAE 635 TABLE I. (continued) Species v M Jussiaeeae Ludwigia 33e odds (Jacq.) Raven, Raven 6571 0.73 313 L. octovalvis, Raven 18670 0.67 287 L. 5 nsis (Cambess. ) Hara 1.54 548 Epilobieae Epilobium colchicum Alboff ee i dn um 1.38 242 2. paniculatum Nutt. ex Torr. & ly 0.52 122 E. (Zauschneria) canum subsp. canum 0.11 17 Hauveae Hauya elegans DC. subsp. cornuta (Hemsley) Raven & Breedlove, we 6432 3.20 1,328 H. elegans subsp. cornuta, Breedlove 10589 3.38 1,153 H. elegans subsp. co ornuta, ~ rlquist VI-1958 2.96 1,270 Hauya heydeana Donnell Smith 1.43 688 outside 3.50 1,771 Hauyeae combined: 2.89 1,242 ceous species and species groups. More recently (Carlquist, 1977; Carlquist & DeBuhr, 1977) I have constructed additional indices for wood ecology. One of these, which may be termed “vulnerability” (V in Table 1) consists of the mean vessel diameter divided by the mean number of vessels per mm? of transection. Although vessel diameter and number of vessels per mm? are roughly inversely proportional for mesophytes (Carlquist, 1975b: 183), xerophytes tend to have narrower vessels and more numerous vessels per mm? than would be expected on the basis of study of mesomorphic woods only, as shown by the desert shrubs included in the graph just cited. This “redundancy” of vessels in xerophytes pro- vides a conductive tissue in which a large number of vessels could be disabled by air embolisms without appreciably lessening conductive capacity. Thus a low value for “vulnerability” can be construed as a high degree of “safety” under water stress conditions and, therefore, xeromorphy. Vessel element length seems a sensitive indication of xeromorphy or meso- morphy (Carlquist, 1975b). Although one could argue that vessel diameter and number of vessels per mm? are somewhat interrelated, vessel element length is controlled entirely independently, by length of fusiform cambial initials. Long vessel elements can be hypothesized to occur in mesomorphic conductive sys- tems, short ones in xeromorphic woods. Therefore, multiplying the “vulnerabil- ity” index by mean vessel element length yields a figure which is termed “meso- morphy” (M in Table 1) here. The higher the value of this value, the greater the hypothetical mesomorphy of the wood. The family Penaeaceae illustrated that this index is reliable on the basis of known species (Carlquist & DeBuhr, 1977). It also appears to be reliable in Onagraceae. Because Onagraceae occupy a notably wide range of ecological situations, I am presenting here the two indi- ces for all species of Onagraceae studied to date (Table 1). One can interpret these indices only within ranges. A “V” figure markedly 636 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 below 1.0 (perhaps 0.1-0.5) would indicate a high degree of xeromorphy ( Carl- quist, 1977), and the lower limit for true mesomorphy would be close to 3.0. With respect to the "M" index, true xeromorphy is indicated probably by a level below 30.0. In Tremandraceae, values range between 15.4 and 45.0 (Carl- quist, 1977), and the highest “M” value in Penaeaceae was 587 (Carlquist & DeBuhr, 1977). Small sampling differences can alter values of these indices, obviously. Future development of these indices will indicate their potential validity in ecological and physiological analyses of dicotyledon families. How- ever, the V and M indices show the same sequence (Hauyeae; Fuchsieae; large shrubby Lopezieae; annual or suffrutescent Lopezieae; annual Onagreae; peren- nial Onagreae) as was demonstrated by the vessel diameter + vessel element length figure (Carlquist, 1975a). The latter figure, however, was higher for the Jussieeae than for Hauyeae, a fact I related to the hydrophytic habit of Jussieeae. However, the V value for Jussieeae is low, which one might not expect in a hydrophyte. Perhaps the fact that Jussieeae grow in seasonally wet but often drying environments explains why they have xylem that combines low vulnera- bility with mesomorphy. A relatively high amount of redundancy in vessels in Jussieeae would cope well with the drying out of habitats. The high V value for Hauyeae (and, to a lesser extent, Fuchsieae) would correlate with lack of any marked water stress in the habitats of these species. If one can interpret the M value for inside wood versus outer wood in a log, Fuchsia excorticata and both species of Hauya begin with appreciably less meso- morphic wood structure and increase in mesomorphy, as well as vulnerability, with age. This would not be unexpected in view of my earlier (1975b) consid- erations. However, if this phenomenon does prove to be widespread in dicotyle- dons, we must conclude that vascular plants tend to have less vulnerability, less mesomorphy in earlier-formed xylem. If the root system of a woody dicotyledon would be expected to experience more severe fluctuation in water availability (nearer the soil surface) than when a tree is older, this would be logical. How- ever, study of additional species would be valuable in this respect. Future application of these indices will indicate their potential validity in physiological analyses of dicotyledon families. Onagraceae is, however, a critical family in this respect, and appears to reflect the broad range of ecology of the family as a whole, and the ecology of individual species and of growth forms, as discussed earlier (1975a) and here. LITERATURE CITED Ball, I. W., & W. W. Tupper. 1918. Size variation in tracheary cells. I. A comparison between us secondary xylems of vascular cryptogams, gymnosperms, and angiosperms. Proc. Amer. Acad. Arts 54: 149-204. BUTTERFIELD, B. G. & . MEYLAN. 1973. Scanning electron micrographs of New Zealand woods. 3. Fuchsia excorticata (J. R. & G. Forst.) Linn. f. New Zealand J. Bot. 11: 411- 41 CanLQuisr, S. 1975a. Wood anatomy of Onagraceae, with notes on alternative modes of photosynthate movement in dicotyledon woods. Ann. Missouri Bot. Gard. 62: 386-424. 1975b 5l Ecological Strategies of Xylem Evolution. Univ. of California Press, Berkeley. 1977. Wood anatomy of Tremandraceae: phylogenetic and ecological interpreta- tions, Amer. J. Bot. 64: 704-713. 1977] CARLQUIST—WOOD ANATOMY OF ONAGRACEAE 637 L. DeBunr. 1977. Wood anatomy of ae (Myrtales): comparative, phy- ie and ecológica] implications. Bot. J. L Soc. (in press). Moss, E. H. 1936. The ecology of Epilobium paid sik der with particular reference to rings of periderm in the wood. Amer. J. Bot. 23: 114-120. A NEW SPECIES OF LOPEZIA (ONAGRACEAE) FROM SINALOA, MEXICO' PETER H. RAvEN? ABSTRACT Lopezia concinna Raven is described from Sinaloa, Mexico. It is unique in the genus in its elaborately . peii and wide lower sepals. It appears to be closely related to L. conjungens. See Within the largely Mexican genus Lopezia, the evolution of distinctive annual taxa at the relatively arid margins of the range is a pronounced trend (Plitmann et al., 1973; Plitmann et al., 1975). Within sect. Lopezia, a group of nine species, three such annual derivatives—L. cornuta S. Wats., L. ciliatula Plitmann, Raven & Breedlove, and L. conjungens T. S. Brandegee—occur within the state of Sinaloa, and the latter two are restricted to it. Nevertheless, the discovery of a distinctive and very handsome novelty within this group by James L. Reveal and Raymond M. Harley, to whom I am most grateful for the privilege of study- ing their material, is of considerable interest, and suggests the possibility of further additions to this genus, which now totals 22 species. Lopezia (sect. Lopezia) concinna Raven, sp. nov.—Fics. 1-2. Species sepalis dimorphis 10-13 mm longis, petalis superioribus 14-16 mm longis ornate notatis eglandulosis ab aliis sectionis Lopeziae diversa. Erect annual. Stems 3-7 dm tall, well branched, terete, reddish, hirsute with white hairs. Leaves 2.5-5.7 cm long, 1.4-2.5 cm wide, ovate, truncate or rounded at the base, acute or acuminate at the apex, subentire or shallowly serrulate- crenulate, subglabrous or with a few scattered hairs along the midrib below and along the margins, with 5-8 veins on each side of the midrib, mostly subop- posite; petioles 8-30 cm long, hirsute. Pedicels 10-14 mm long, spreading, gla- brous. Sepals 10-13 mm long, the upper three 1-1.5 mm wide, linear and keeled at the apex, the single lower one 3-3.5 mm wide and lanceolate, keeled along its entire length. Lower petals 11-14 mm long, 6-7.5 mm wide, subovate, entire, clawed, the claw 2-3 mm long, the petals entirely fuchsia purple with a dark spot at junction of the claw; upper petals 14-16 mm long, 1.5-2 mm wide, linear, shortly clawed, subacute at the apex, slightly auriculate at the base, fuchsia purple with a narrow V-shaped dark stripe, a parallel white stripe, then a thick parallel orange stripe, a broad parallel white stripe, and a final ornate dark stripe in the third of the limb just about the auricles; glands absent, but the flower evidently nectariferous. Fertile stamen 8-9.3 mm long; filament com- Support from the U. S. National Science Foundation is gratefully acknowledged. Peter Hoch provided valuable technical assistance, Steven R. Seavey the chromosomal information, and Richar Eyde comments on the floral anatomy. The herbarium of the University of California, Berkeley (UC), kindly lend the type and only known specimen of Lopezia conjungens. Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110, U.S.A. ANN. Missouni Bor. Garp. 64: 638-641. 1977. 1977] RAVEN—LOPEZIA 639 Ficures 1-2. Flowers of Lopezia concinna Raven, x 2.7; photographs by Robert Srenco. —1. Before explosive release of the fertile stamen by the staminode. Note drops of nectar at the base of the upper petals.—2. After release of the fertile stamen. The staminode has curved downward, the Fertile: stamen upward, and the style elongated, carrying the stigma into the position formerly occupied by the staminode holding the fertile stamen under tension. pressed, 6-7 mm long, 0.8-1.2 mm wide, dark red at the base, shading to pink at the point of anther attachment; anther 2-2.5 mm long, 0.2-0.3 mm wide, ruby red. Staminode 8-9.5 mm long, suborbicular or obovate-spatulate, retuse at the apex, abruptly narrowed to a claw 44.5 mm long, ruby red, the claw pink. Style 8-10 mm long, pink; stigma 1.4-1.6 mm in diameter, white. Ovary broadly ellipsoid to rotund, 2.5-4 mm long, 2-3.2 mm wide, glabrous. Capsule 6-7 mm long, 3.54.5 mm thick, obovoid-elongate, fleshy. Seeds several in each locule, 1.3-1.6 mm long, oblong-ovoid, uncurved, with transverse protuberances over the entire surface, brown. Gametic chromosome number, n — 10 (10 bivalents at meiotic metaphase I). Tyre: Mexico. sINALOA: Along the dirt road from Rosario to Plomosas, about 3.5 km east of La Rastra and 3.2 km up the grade from a river crossing, this about 1.5 km south of Rosario, on a rocky road cut along the steep canyon wall, 8 Oct. 1975, James L. Reveal & Raymond M. Harley 4064 (MO-2412198, holotype; CAS, IPN, K, MEXU, MICH, US, isotypes). Distribution: Known only from the type collection. See p. 641. This elegant new species, with its beautiful flowers (Figs. 1-2), differs from all other members of its section, except L. conjungens, in its lack of glands on the upper petals. That species is also known only from its type collection, and we have been unable to examine living material. Its type and only known locality is some 270 km northwest of the locality in southernmost Sinaloa where L. concinna was discovered. The only known plants of L. conjungens are sub- glabrous, have much smaller and paler flowers, and lack the elaborate markings of the petals of L. concinna. Indeed, the markings in the petals of L. concinna and its conspicuously wider lower sepal are absolutely distinctive within the 640 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1977] RAVEN—LOPEZIA 641 genus. Its flowers are substantially larger than those of all other species of the section except for those of the very different L. suffrutescens Munz. The seeds of the two species (Figs. 3-8) both have transverse ridges, but those of L. con- cinna are much coarser and occupy the entire surface of the seed, whereas those of L. conjungens are fine and widely spaced. The epidermal cells on the seeds of L. conjungens are oblong, while those on the seeds of L. concinna more nearly square. Finally, the seeds of L. conjungens are more markedly incurved than those of L. concinna. More material of each species, and especially living mate- rial of L. conjungens, will be necessary to clarify their relationships, which ap- pear close; further, the two species may well have been derived from a common ancestor, or L. concinna may have given rise to L. conjungens. We have not observed nectar production, which is copious in L. concinna and apparently arises from the base of the petals, in any other species of sect. Lopezia. Richard H. Eyde sectioned floral material of L. concinna, from prog- eny of the type grown at the Missouri Botanical Garden, and found the order of divergence of the parts to be as reported for L. hintonii Foster [= L. miniata Lag. ex DC. subsp. hintonii (Foster) Plitmann, Raven & Breedlove] by Eyde & Morgan (1973), with nectaries in the usual position for Lopezia. The chromosome number was determined in progeny of the type collection grown in the experimental greenhouse at the Missouri Botanical Garden. Details of the flowers have also been studied in this cultivated material. At its type and only known locality, L. concinna was relatively common in protected, dripping wet recesses along a north-facing cliff-face in a forest domi- nated by trees of Bursera with Hyptis, Salvia, Polymnia, Lasiacis, Malvaviscus, and Euphorbia subg. Poinsettia common in the understory. Directly associated with the plants of the Lopezia were Amemia affinis Baker, Cuphea llavea Lex., Peperomia sp., Pinguicula crenatiloba DC., Polypodium pumila ( Bonpl.) Cogn., Pterolepis pumila (Bonpl.) Cogn., and Salvia misella Kunt LITERATURE CITED Eype, R. H. & J. T. Morcan. 1973. Floral structure and evolution in Lopezieae ( Onagra- ceae ). aa J. Bot. 60: 771-787. PLIrMANN, U., P. H. Raven & D. E. BREEDLOVE. 1973. The systematics of Lopezieae re Ann. Missouri Bot. Gard, 60; 478-563. „W. Tat & D. E. BREEDLOVE. 1975. Cytological studies in Lopezieae ( Ona- anes | Bot. Gaz. (Crawfordsville) 136: 322-332. NOTE ADDED IN PROOF While this article was in press, D. E. Breedlove kindly sent a second collection of the new species from 80 km farther east: Durango, canyon of Rio Mezquital near Nayarit, ca. 750 m, 1-6 Nov. 1977, G. H. Bolton 101 (CAS, MO < ‘GuREs 3-8. Scanning electron micrographs of seeds of two closely related species of Lopezia sect. Lopezia. All from the respective type 5 The bars at the top of Ex plate indicate, respectively, 0.5 wm, 125 um, and 25 um.—3-5. L. conjungens.—6-8. concinna. REINTERPRETATION OF THE TYPE OF GODETIA BOTTAE SPACH (ONAGRACEAE)! PETER H. RAVEN? AND DENNISs R. PARNELL? Paolo Emilio Botta, who traveled in the Héros, commanded by August Ber- nard Duhaut-Cilly, collected birds and plants along the coast of California in 1827 and 1828 (McKelvey, 1955). One of these was the type of the taxon later described as Godetia bottae Spach (1835). It is preserved in the Herbarium of the Muséum National d'Histoire Naturelle, Paris (P), and is annotated in Edouard Spach’s hand, “Godetia Bottae, nob. (Spach, 1839), California, M. Botta.” A second specimen is annotated simply “Godetia bottae” with an inscrip- tion in another hand that says "Monterey, M. P. E. Botta, 1829.” According to Dr. A. Lourteig, this second hand is that of Botta. At any rate, these specimens are not the species that has subsequently come to be known as Clarkia bottae (Spach) Lewis & Lewis (1955), but rather the one known as C. deflexa (Jeps.) Lewis & Lewis. Harlan Lewis concurs in this redetermination. Judged from the ports that the Héros visited regularly and the time of year, they were probably collected in the summer of 1827 or that of 1828 either at Santa Barbara or at San Pedro; the species does not occur at or near Monterey and the notation on the second herbarium specimen must have been made in error. Collections from Monterey County assigned by Lewis & Lewis (1955) to the taxon C. deflexa have been shown to comprise a distinct species described as C. jolonensis Parnell (1970). In view of these findings, the following new synonymy is appropriate: Clarkia (subsectio Peripetasma Lewis & Lewis) lewisii Raven & Parnell, sp. nov. Clarkia bottae sensu Lewis & Lewis, Univ. Calif. Publ. Bot. 20: 315. 1955; non Godetia bottae Spach, Nouv. Ann. Mus. Hist. Nat. 4: 393. 1835. C. cylindrico differt: tubo floralis annulo pilorum summo intus, 1.5-3 mm longus, non colorato intus. Type: U.S.A. CALIFORNIA: Monterey Co., Point Lobos, along the trail to China Cove from the end of the road, 26 June 1947, H. and M. Lewis 498 (LA). This species is dedicated to Harlan Lewis of the University of California, Los Angeles, who has made Clarkia one of the groups that has contributed most to our understanding of plant evolution. As two of his former graduate students, we feel a sincere debt of gratitude to him. ! Support from the U. S. National Science Foundation through grants to Peter Raven is gratefully acknowledged. Dr. Lourteig iun Professor Harlan Lewis have helped us in the er ee ition of the specimens e PSA d he Missouri Botanical Garden, 2345 Tower G rove Avenue, St. Louis, Missouri 63110. ` Department of Biological Sciences, California State Universite at Hayward, Hayward, California 94542. ANN. Missovuni Bor. Garp. 64: 642-643. 1977. 1977] RAVEN & PARNELL—GODETIA BOTTAE 643 Clarkia bottae (Spach) Lewis & Lewis, Madroño 12: 33. 1953. Godetia bottae Spach, Nouv. Ann. Mus. Hist. Nat. 4: 393. 1835. Oenothera bottae (Spach) Torr. & A. Gray, Fl. N. Amer. 1: 505. 1840. Godetia bottae Spach var. deflexa (Jeps. ) Hitche., Bot. Gaz, (Crawfordsville) 89: 355. 1930. Clarkia Qo (Jeps.) Lewis & Lewis, Madroño 12: 33. 1953; Lewis & Lewis, Univ. Calif. Publ. . 20: 334. 1955. With the description of Clarkia jolonensis Parnell (1970) and C. rostrata Davis (1970), the number of species of sect. Peripetasma Lewis & Lewis is now 11. LITERATURE CITED Davis, W. S. 1970. The systematics of Clarkia bottae, C. cylindrica, and a related new species, C. rostrata. Brittonia 22: 270-2 Lewis, H. & M. E. Lewis. 1955. The genus C Clarkia. Univ. Calif. Publ. Bot. 20: 241-392. McKELvEy, S. D. 1955. Botanical Exploration of the Trans-Mississippi West 1790-1850. Arnold Arboretum of Harvard Univ., Jamaica Plain, Mass. xl PARNELL, D. R. 1970. Clarkia jolonensis (Onagraceae), a new species from the inner Coast Ranges of California. Madroñ Seac, É. 1835. Monographia 5 ‘None Ann. Mus. Hist. Nat. 4: 320-408, pl. 30-31. REPRODUCTIVE STRUCTURES AND EVOLUTION IN LUDWIGIA (ONAGRACEAE). L ANDROECIUM, PLACENTATION, MERISM' RICHARD H. EYDE” ABSTRACT This article, based on serial sections from 19 species of Ludwigia (supplemented where necessary with preparations from other Onagraceae), begins an effort to outline the evolution of flower, fruit, and seed characters within the genus and to link the outline with what is known of floral evolution elsewhere in the family. New í include the following: all Lud- wigia anthers have a prominent endothecium, and developing anthers of certain advanced spe- cies are markedly H-shaped in cross section; pollen of two species matures in isolated packets; ovules of L. leptocarpa, though commonly l-seriate, can be distally pluriseriate; only rarely does a Ludwigia placenta have a median groove suggesting paired carpel margins. The deeply intrusive placentas seen in section Myrtocarpus, but lacking in some of the other sections, are probably ancestral, and the old idea that diplostemony and 4+-mery are ancestral holds up well when reexamined critically. Few families have been as intensively studied by evolutionary botanists as the Onagraceae. Relationships among many infraspecific variants and among closely connected species groups have been firmly established through cytologi- cal work, breeding experiments, and field observations of reproductive events. As one proceeds to more widely separated taxa, however, biosystematic methods become inapplicable; consequently, evolutionary links among the genera of Ona- graceae are not yet well understood. Structural comparison remains the best— perhaps the only—way to improve our knowledge of these links. First, structural differences among the taxa must be identified, then the direction of evolutionary change can be inferred by critically weighing the alternatives. The Onagraceae are ideal in several respects for comparative studies of floral structure. For one thing, the family is of manageable size: Raven currently rec- ognizes 17 genera and estimates the number of species to be 600-700. “Spirit collections” of many of these species are available for anatomical work because of the research efforts of Raven and his collaborators. Another advantage in working with Onagraceae is that the taxa are diverse enough to be challenging, yet undoubtedly of common evolutionary origin. Among the characters that show the Onagraceae to be a natural family are the peculiar viscin threads on onagra- ceous pollen (Skvarla et al., 1977) and the distinctive 4-nucleate embryo sac (Seshavataram, 1970; Bhatnagar & Johri, 1972:91; Palser, 1975:641). Still an- other advantage is that the nearest extra-familial affinities of the Onagraceae a known to be among the myrtalean families Combretaceae, Crypteroniaceae, Lythraceae, Melastomataceae, Myrtaceae, Punicaceae, and Sonneratiaceae. Sim- ilarities in floral structure within this alliance were recognized by pre-Darwinian taxonomists and are now seen as indicators of shared ancestry, with strong con- = c 11 thank P. Raven and T. P. Ramamoorthy for criticizing the typescript. The National Science Foundation contributed indirectly, via a series of grants to Raven, by supporting the Smithsonian photographer V. Krantz and museum specialist S. Yankowski, respectively. * Department of Botany, Smithsonian Institution, Washington, D.C. 20560. ANN. Missouni Bor. GARD. 64: 644-655. 1977. 1977] EYDE—EVOLUTION IN LUDWIGIA 645 firmatory evidence from such diverse sources as embryology (Subramanyam, 1951) and vegetative anatomy (Carlquist, 1975; van Vliet, 1975; van Vliet & Baas, 1975). Ideas on “ancestral versus derived” in the Onagraceae can be tested by looking into the related families for satisfactory distribution of the putative an- cestral state. Though the ultimate goal is to understand the evolution of the Onagraceae as a whole, this report concentrates on Ludwigia L., the only genus of the tribe Jussiaeeae (Raven, 1963). More than 70 species are known, all from wet habitats in temperate and tropical regions around the world (for illustrations, see Micheli, 1875; Rickett & Collaborators, 1967; Correll & Correll, 1972). Various authors have considered Ludwigia—or Jussiaea, now a synonym of Ludwigia—the primi- tive onagraceous genus (see Melchior, 1964; Takhtajan, 1966, 1973) because it seemed to provide the best link with adjoining families and because Ludwigia flowers were thought to retain ancestral traits, among them the 5(or more )- merous condition and the absence of a floral tube beyond the inferior ovary. It now seems that the absence of a floral tube is secondary in this case; moreover, it is now recognized that Ludwigia has a derived basic chromosome number and other specialized features. Undoubtedly, however, the genus represents an early evolutionary offshoot within the Onagraceae; a credible phylogenetic outline of the family must account for its peculiarities. My wet material of Ludwigia, flowers from 30 collections belonging to 19 species, is listed in Table 1 along with nine collections from five more species (asterisked ) that were available only as herbarium specimens. Altogether, these collections represent 10 of the 17 infrageneric sections recognized by Raven in 1963. Stained serial cross sections were prepared from all the collections and replicate series from most, also longitudinal series as needed, bringing to more than 100 the number of flowers (and developing fruits) sectioned and examined. Thanks to Raven and his collectors, I was able to compare the sectioned Lud- wigia flowers with sectioned flowers from more than 70 additional species of Onagraceae, systematically selected from all parts of the family. I begin with the androecium, though I have few new observations on Lud- wigia stamens, because I want an unequivocal basis for discussing character associations, and I think all systematic botanists, despite differences in training and philosophical outlook, will accept the evidence for ancestral diplostemony in Ludwigia. ANDROECIUM In general, Ludwigia species are constantly diplostemonous (the old genus Jussiaea, Fig. 1) or constantly haplostemonous (Fig. 2), but L. perennis can be intermediate (Raven, 1963), and at least one species, L. inclinata, includes some plants with two whorls of stamens, others with one whorl (Raven, personal com- munication). One may be confident that diplostemony is the ancestral condition because of its wide distribution in the Onagraceae, haplostemony occurring (out- side Ludwigia) only in two genera with highly specialized flowers (Circaea, Lopezia) and in specialized members of two other genera (one species of Camis- sonia, sect. Eucharidium of Clarkia). In neighboring families, stamens are 646 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE l. Herbarium vouchers for sectioned Ludwigia flowers.“ Taxa. Vouchers Sack - Oligospermum eploides (H.B.K.) Raven Raven 8 (LA), California. Raven 26493 (MO), rkans: L. uruguayensis (Camb.) Hara Raven s.n. (Ds). California [Naturalized, Stanford Univ. ]. Sect. Oocarpon L. torulosa (Arnott) Hara Cowan 38886 (US), French Guiana. de la Cruz 3813 Guyana. Howard & Howard 9911 (US), Dominican Republic Sect. Nipponia 2E; ee Maxim. Chien 207 (US), China. Sect. Seminuc L: 1 (Nutt.) Hara Chevalier 21 (DUKE), Florida. Raven 26491 (MO), rkansas. Sect. Fissendocarpa *L. hyssopifolia (G. Don) Exell Asplund 14132 (US), Peru. Purpus 6973 (US), Mexico. Sect. 85 yrtocarpus d E Walt. idus Mr 896 (both DUKE), North Carolina. 1 26469 (MO), Arkansas. L. densiflora ~ ) Hara Ma 3940 (US), Brazil. L. erecta (L.) Har Raven 21573 (DS), Costa Rica. L. foliobracteolata (Munz) Hara Raven 21981 D Costa Rica L. latifolia ( Benth.) H ie ven 21575 (DS), Costa Rica L. peruviana (L.) His Steinberg s.n. (FAU), Florida. *L. tomentosa (Camb.) Hara Dawson 15154 (RSA), Brazil. Gardner 2571 (US), Brazil. Sect. Macrocar] L. 5 (Munz) Hara Krapovickas & Cristóbal 12089 (DS), Paraguay. L. octovalvis (Jacq.) Raven Raven 21574 (DS), Costa Rica. Sect. Ludwigia . alternifolia L. Broome 851, 860, 862 (all DUKE), North Carolina. L. maritima Harper Chevalier 18 (DUKE), Florida. Arguelles 1 (MO), ississippi. L. virgata Michx. Broome 863 (D 3 s Carolina. Willingham 597 (MO), Georg Sect. Microcarpium : Ell. Arguelles 3 (MO), Mississippi. L. glandulosa Walt. Broome 865 (DUKE), North Carolina. L. lineari: al Broome 856 (DUKE), North Caroli L. pilosa Walt. Broome 861, 902 (both DUKE), North Carolina. Sect. Dantia L. arcuata Walt. Chevalier 11 (DUKE), North Carolina. L. palustris (L.) Ell. Broome 859 (DUKE), North Carolina. Willingham MO), Georgia. Arguelles 2 (MO), Mis- sissippi. * Asterisks mark species for which herbarium flowers were sectioned; for other species, liquid- preserved Haw were used. commonly twice or more than twice the number of sepals, the haplostemonous exceptions being the apetalous genus Crypteronia, the myrtaceous genus Myr- rhinium (with specialized inflorescences and stamens; McVaugh, 1968: 407), and certain members of the families Lythraceae and Melastomataceae. The an- droecia of Lythraceae are almost bewildering in their meristic diversity; how- 1977] EYDE—EVOLUTION IN LUDWIGIA 647 L d 1a 2b Ficures 1-2. — illustrations —1l. L. leptocarpa. Top of plant (a). x 0.3. Flower (b). x 1.7 5 L. linearis. Top of plant (a). x 0.3. Flower (b) and fruit (c). x 3.3. Partly redraw Aq bç . Tangerini ps oe prepared by G. Reinert for R. K. Godfrey, who kindly eat ic 3 for copyi ever, the fact that some of the haplostemonous taxa have stamens opposite the petals and others have stamens opposite the sepals is best explained by deriving both forms from precursors with at least two whorls of stamens. In the Melasto- mataceae, haplostemony is very much a minority trait, but a widely scattered one, occurring in seven New World genera (Wurdack, 1971: 360) and at least 648 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 five Old World genera: Blastus, Dactylocladus, Dissochaeta, Omphalopus, and Sonerila. I am assured by melastome specialist J. Wurdack that this taxonomic distribution indicates multiple evolutionary derivation of haplostemony within the family. Returning to the Onagraceae, we find in the genus Ludwigia itself that the woodier tropical species are mostly diplostemonous, whereas temperate species with such advanced features as poricidal capsules or apetaly are haplo- stemonous, still another indication that diplostemony is ancestral. The stamens of Ludwigia species are much alike externally except for those of sect. Ludwigia. In this section, the filament joins the versatile anther in a deep dorsal groove, and the halves of the anther are parallel during development; consequently, the cross section is decidedly H-shaped, unlike that of a develop- ing anther in other sections of the genus (cf. Figs. 3-4). All the examined ludwigias have a conspicuous endothecium or “fibrous” layer. That is, the hypodermis of the anther is a layer of relatively large cells with narrow wall thickenings. As regards the development of the endothecium, Ludwigia anthers are like anthers of Circaea, Hauya, and certain fuchsias (e.g., F. arborescens); they are unlike anthers of Clarkia, Gaura, and Gayophytum, in which the endothecial cells are notably smaller than the epidermal cells. Future studies of onagraceous flowers should take careful note of the anther wall, for there are clear-cut endothecial differences not only among the genera but also within certain genera ( Epilobium, Fuchsia). A prominent endothecium is prob- ably ancestral (see Eames, 1961: 138 ff.); so these differences could turn out to be valuable evolutionary clues. In my material of Ludwigia latifolia and of L. linearis, the developing pollen grains are in packets that are separated from other packets above and below by bands of parenchyma. This observation is of more than passing interest because interrupted sporogenous tissue was known heretofore in only five onagraceous genera—Hauya and four genera of the tribe Onagreae—and Raven (1969: 161) has argued, contra Munz (1965), that the shared character makes Hauya a member of the Onagreae. Discovery of pollen packets in another tribe, where they appear to have evolved twice, undermines Raven’s argument and makes Hauya's placement problematic. PLACENTATION In certain Ludwigia species, notably those of sect. Oligospermum and Semi- nuda, the ovules are inserted in l-seriate rows, one vertical row to each locule (Figs. 5-6). In sect. Ludwigia, Macrocarpon, and Myrtocarpus, pluriseriate ovules are crowded on deeply intrusive placentas that are commonly spatulate in cross-section (Fig. 8). Sections Dantia and Microcarpium also have pluriseri- ate ovules, but the placentas are not spatulate in cross section (Fig. 9). In L. epilobioides, the ovules are l-seriate in most locules; they can also be more or less 2-seriate (irregularly so), and both arrangements can occur in the same ovary. Ludwigia hyssopifolia is unusual in that the ovules are irregularly pluri- seriate at the distal end of the placentas and l-seriate below; I have observed the same situation in one collection of L. leptocarpa, sect. Seminuda (Fig. 7). In all its variations, Ludwigia placentation has advanced beyond that of most 1977] EYDE—EVOLUTION IN LUDWIGIA 649 Ficures 3-8. Laduiala flowers in cross sectio shaped 1 of anther. x 80. L. decurrens, ree 855. Con In both figs. the abaxial side of the anther is up.—5. i —7. L. leptocarpa, Chevalier 21. Here the upper part of the ovary contains pluriseriate ovules. x 20.—8. L. foliobracteolata, Raven 21981. Note deeply sop HO eA x 17. 650 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Onagraceae, for one rarely finds a vestige of the ancestral bipartite placental structure, and an actual separation of the ovarian septa (opening of the ventral sutures) can be found only by following them upward into the style (Fig. 11). This reasoning may seem uncritically “classical” to some readers; so I shall take pains to explain. If we begin with the formalistic concept of carpel closure endorsed by Eames and others before him, then visualize the divisions of the onagraceous ovary as imbedded carpels and the placentas as fused carpel margins, we can say of Ludwigia that "the fusion is anatomically complete and the placenta is simple in form and structure" (Eames, 1961: 205). To be sure, evolutionary morpholo- gists now find Eames's interpretation of syncarpy inadequate because it does not take into account a change in gynoecial ontogeny that has occurred within many groups, including the Myrtales—namely, a shift in the locus of septal develop- ment and ovule inception from discrete carpel primordia to a more recently evolved tubular meristem (the gynoecial cylinder of Sattler, 1973) beneath the carpel primordia. And associated with the origin of the Onagraceae there has been a further modification of the ontogenetic sequence so that the ovules now develop on septa growing upward and inward from a meristematic tube under- lying the gynoecial cylinder as well as all the floral primordia (Sattler, 1973). It is not altogether wrong, however, to view the angles formed by the ovarian septa as fused carpel margins, at least in the upper part of the ovary where the septa are ingrowths that actually unite as the ovary develops. After all, the septa do not arise in random positions. They are initiated in line with the ends of the crescent-shaped carpel primordia above them, presumably under the morphogenetic influence of the carpel primordia? Moreover, when the devel- oping onagraceous flower first produces septa, then placentas, then ovules within its inferior ovary, it repeats a canalized sequence that began in distant ancestors with superior, apocarpous gynoecia. One cannot argue otherwise, I think, with- out opting for polyphylesis of angiosperms. As the gynoecium changed from apo- carpus to syncarpous and from superior to inferior, the placentas continued to develop from the inner portions of the septa, and the septa continued to develop in vertical alignment with the margins of the increasingly ephemeral carpel pri- mordia. (In Ludwigia, carpel primordia persist only as obscure stigmatic lobes.) From the evolutionary standpoint, therefore, the upper part of the ovary in many Onagraceae does contain carpel margins, even though they are no longer direct outgrowths from the carpel primordia, and the degree to which these margins In Lythrum salicaria (Lythraceae ), where the two carpel spout appear fleetingly, if at all, the location of the ovary’s partition is not fixed. Within a single orescence, one may find some ovaries divided in de median plane, others in the tranversal Sine (Sattler, 1973). -> Fıcures 9-14. Onagraceous flowers in cross section.—9. Ludwigia palustris, Willingham 598. Plac i ied do not have the swollen appearance of those in Fig. 8. x 52.— 10. Hauya ele- gans, Breedlove 6432. Ovary of a 5-merous flower showing discrete od margins. x 12.— 2 * Raven 21574. Base of style showing separation of septa; a stamen diverges at right. x 45.—12. Epilobium fleischeri, cultivated at the Royal Botanic Garden, Edinburgh, EYDE—EVOLUTION IN LUDWIGIA à Ey wr Wes ` 2 B H H - they PET 4 ma : sat re ue ‘ "à - pag segs? as C8092. Uppermost level of ovary showing separation of septa. X 53.—13. section taken about 600 u below Fig. 12 showing median placental grooves. peruviana, Steinberg s. n. grooves (arrows) in two of the placentas. X £ Same [»] x 53.— flower, I4. E. aevel of divergence of floral appendages showing distinct median 652 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 are fused can be an indication of evolutionary advancement. I have argued else- where that the union of carpel margins has reversed to some extent in certain Rosaceae (Eyde, 1975). For angiosperms as a whole, however, I accept the gen- eralization that fused margins are derived and unfused margins ancestral. To test the applicability of this generalization to the Onagraceae, we can compare gynoecia of the woody tropical genus Hauya with gynoecia of Epilo- bium, a predominantly herbaceous genus of temperate and cold regions. If we were to take serial cross sections of a Hauya flower and project them rapidly on a screen, proceeding from base to apex, we would see the septa separate within the ovule-bearing region (Fig. 10). If the sections were from a fully developed flower, the radii along which the septa part would be marked by pollen-trans- mitting tissue. If the projected sections were from a flower of Fuchsia, another woody tropical genus, septal separation would also be observed within the fertile part of the ovary. But a similar sequence through an Epilobium flower would differ in that the septa would not separate, if at all, until we had gone beyond the ovules into the very summit of the ovary (Fig. 12). The ontogenetic expla- nation for this difference is that the zone of septal upgrowth is relatively greater in Epilobium and the zone of septal ingrowth relatively less (Kaienburg, 1950: 400). Despite the fact that the zone of septal ingrowth is confined to the upper- most part of the ovary in the finished flower of Epilobium, each plane of septal fusion can be followed downward through much of the ovary because its posi- tion is marked by a median groove in the placenta (Fig. 13). Similar placental grooves occur in most onagrads, even those with only one ovule per locule, though they are not always as distinct as they are in Epilobium. The taxa I have found to be exceptional—that is, lacking a well-marked placental groove—are Circaea, two species of Oenothera (O. campylocalyx, O. rosea), and most species of Ludwigia. In some species of Ludwigia, such as L. hyssopifolia, an observer passing through the gynoecium from base to apex might enter the style before seeing the separation of the septa, but in certain species of sect. Myrtocarpus and its derivative sect. Ludwigia there is an “elevated disc" below the style (see Micheli, 1875, for illustrations) in which the parting of the septa can be observed. Fur- thermore, it is only in species belonging to these two sections (L. latifolia, L peruviana, L. virgata) that I have seen any trace of a placental groove, and then only in the upper part of the ovary (Fig. 14). If my reasoning with regard to ancestral and derived placental characters is correct, these observations place sect. Myrtocarpus near the ancestry of the genus, though its deeply intrusive placentas are advanced over those of most Onagraceae. If we test the argument by considering the taxonomic distribution of bipar- tite placentas and partially unfused margins in related families, we find the dis- tribution to be consistent with the view that these features are ancestral. In the Myrtaceae, a family closer than the Onagraceae to the ancestry of the Myrtales, taxa with partially unfused margins within the ovary are found among the cap- sular groups as well as the fleshy-fruited groups (Ludwig, 1952). In the Lythra- ceae, the small tree Lagerstroemia indica has separate septa in the summit of the ovary, whereas the slender herb Lythrum salicaria, a more advanced member 1977] EYDE—EVOLUTION IN LUDWIGIA 653 of the family, has neither separate septa nor grooved placentas (personal obser- vations). Looking into the Sonneratiaceae, Cronquist’s (1968) choice as the most nearly ancestral myrtalean family, we find that the septa are separate in the upper fifth of the Sonneratia ovary (Mahabalé & Deshpande, 1957). To judge from published illustrations, the ovaries of Duabanga, the only other genus of the Sonneratiaceae, are structurally similar (Jayaweera, 1967: figs. 1J, 3F). In the highly specialized flowers of Melastomataceae, however, the septa part in the style (van Heel, 1958) or not at all (Eyde & Teeri, 1967; Subramanyam & Narayana, 1969) DERIVED STATUS OF 4-MERY The evolutionary morphologist of a few decades ago might have claimed derived status for 4-mery without risking contradiction, at least from American colleagues, on the principle that the “polymerous flower structure precedes, and the oligomerous structure follows from it” (Bessey, 1915). Many exceptions to this principle are known (Stebbins, 1967), however, and Huether's experiments on Linanthus (summarized by Stebbins, 1974) have shown how readily the num- ber of floral parts can be increased as well as decreased under selective stress. In Ludwigia, moreover, higher numbers of floral parts can occur in associa- tion with certain advanced features. For instance, in sect. Oocarpon, with 5- merous flowers, and in sect. Oligospermum, where 5-mery is the rule and 6-mery occasional, the higher numbers are linked with 1-seriate ovules and a specialized endocarp. In sect. Seminuda, with 4-7-merous flowers, the ovules are also I- seriate, though the fruits are specialized in a different way. Another example is L. epilobioides (sect. Nipponia), a self-pollinating herb of temperate Asia in which 4-6-mery is associated with haplostemony. Raven (1963) reports that 5-merous flowers can be found, albeit rarely, in L. perennis (sect. Caryophyl- loidea); this species too is commonly haplostemonous. In Myrtocarpus, the “phylogenetically central” section of Ludwigia (Raven, 1963), L. densiflora has 4-6-merous flowers in a spicate (derived) inflorescence, and L. peruviana, in which 5-mery is encountered fairly often, is an aggressive polyploid colonizer. (On the other hand, T. P. Ramamoorthy informs me that 5-mery is the usual condition in the Brazilian shrub L. tomentosa and that he has seen 5-merous flowers in a number of other species belonging to sect. Myrtocarpus. ) Excepting these sections—and sect. Prieuria with mostly 3-merous flowers (Raven, 1963)—4-mery is quite constant in the genus Ludwigia. I have seen no 4+-merous flowers in sect. Dantia, Macrocarpon, or Microcarpium; and in sect. Ludwigia, I have seen only two 5-merous flowers of L. virgata and no other exceptions. Throughout the remainder of the family, 4-mery occurs with similar constancy (though the flowers of a few taxa regularly have fewer than four parts). A minority of Hauya flowers are 5-merous and five stigmatic rays can occur in Oenothera (Cleland, 1972: 6), also in at least one species of Epilobium (E. dodonaei, personal observation), but I do not know that 5-mery or partial 5-mery has ever been observed in Fuchsia, a genus seemingly as close as any to the ancestry of the Onagraceae. In the Onagraceae, therefore, ancestral status might be claimed for 4-mery 654 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 on the grounds that 4+-mery is almost exclusively confined to Ludwigia, where it often accompanies derived characters. This reasoning runs into difficulty, however, when the reconstruction of onagraceous ancestry is extended beyond the family limits, for the ancestral Myrtales surely had more than four floral parts per whorl. To claim 4-merous ancestry for the Onagraceae, one would have to begin with myrtalean ancestors in which floral parts were indefinite in number; then postulate a derived group, ancestral to all Onagraceae, with floral parts stabilized in whorls of four; then further postulate a return to 4+-mery in each of several lines within Ludwigia. An evolutionary scheme incorporating these steps would be less economical than one in which 4-mery is treated as a derived character. Stebbins (1967) has pointed out that the number of floral parts in a whorl is partly dependent on the number of cells in the floral apex at the time the whorl is initiated; so it is not surprising that higher numbers are often found in larger flowers. The relationship between meristem size and numbers of parts may explain Müllers (1870) observation that individual plants of Brazilian lud- wigias (species unspecified) tend to produce 5-merous flowers first, 4-merous flowers later (see also Huether, 1968: 128), but there is no consistent relation- ship between floral size and merism in Ludwigia. Some large-flowered species are constantly 4-merous, whereas L. torulosa, with very small flowers, is con- stantly 5-merous. LITERATURE CITED Bessey, C. E. 1915. The phylogenetic taxonomy of flowering plants. Ann. Missouri Bot. Gard. 2: 109-16 ge BHATNAGAR, S. P. & B. M. Jounr.. 1972. Development of angiosperm seeds. Pp. 77-149, in T. T. e (editor), Seed Biology. Vol. 1. Academic Press, New York. CanLQuisr, S. 1975. Wood anatomy of Onagraceae, with notes on alternative -n of photosynthate p in dicotyledon woods. Ann. Missouri Bot. Gard. 62: 386—424. E. 197 CLELAND, R. E. : vothera: Cytogenetics and Evolution. Academic Press, New York. ConnELL, D. S. & H. B. f iilud 1972. Aquatic and Wetland iu of Southwestern United States. U.S. Environmental Protection Agency, Washin D.C. Reissued in 2, volumes in 1975 by Stanford Univ. Press, Stanford, Calif. Treo di pp. 1175-1199.] Cnowquisr, A. 1968. The Evolution and Classification. of Flowering Plants. Houghton Eames, A. J. 1961. 1 of the Angiosperms. McGraw-Hill, New York. EvpEe, R. H. 1975. The bases E angiosperm phylogeny: floral anatomy. Ann. Missouri Bot. Gard. 62: 521-537. [Issued 1976.] . TEER. 1967. m anatomy of Rhexia virginica ( Melastomataceae). Rho- dora 69: 163-178. Heer, W. A. van. 1958. The pistil of Bertolonia marmorata Naud. (Melastomataceae). Blu mea, Suppl. 4: 144-148 HuETHER, C. A., Ja. 1968. Epen of natural genetic variability E in the pentam- erous corolla constancy in Linanthus androsaceus ssp. m pays cs 60: M. A. 18 JAYAWEERA, D. M. A. 57. The genus Duabanga. J. Arnold Arbor. 48: 89104. KAIEN BURG, A.-L. 1950. Zur Kenntnis der N und der 1 tung bei einigen s. Planta 38: 377—430. Lupwic, A. 1952. r Fruchtknotenbau d Myrtaceen und seine Bedeutung für die Glied- ME dieser s Inaug.-Diss., Ludwig-Maximilians-Univ., München. [Unpublished; Library of C ongress has a microfilm of the typescript.] MananALÉ, T. S. & J. V. DESHPANDE. 1957. The genus Sonneratia and its fossil allies. Pa- lacobotanist 6: 51-64, pl. 1—4. [Issued 1959.] McVat . 1968. The genera of American Myrtaceae 35 L " 8. an interim report. Taxon 17: 1977] EYDE—EVOLUTION IN LUDWIGIA 655 Metcuior, H. 1964. Reihe Myrtiflorae. Pp. 345-366, in A. Engler’s Syllabus der Pflanzen- familien. M 2. Gebrüder Borntraeger, Berlin. MichELI, M. 1875. Onagraceae. Pp. 146-182, pl. 28-38, in C. F. P. de ue ( succeeded y A. W. ule Flora Brasiliensis. Vol. 13, pars 2. Monachii et Lipsia MürLEn, F. 1870. Botanische Notizen aus einem Briefe von Fritz Miiller. (Ttajahy, den 7. December 1869). Mitgetheilt von F. Hildebrand. Bot. Zeitung (Berlin) 28: 273-275. Munz, P. A. 1965. Onagraceae. N. Amer. Fl., ser. 2, 5: 1-278. Pauser, B. F. 1975. The bases E s atu ‘phylogeny: embryology. Ann. Missouri Bot. Gard. 62: Su [Issued 1 Raven, P. H. 1963. The Old ur species of op 1 Jussiaea), with a syn— opsis of the genus (Onagraceae). Reinwardtia 6: 327 [Issued 1964. 1969. A revision of the genus Camissonia 3 Contrib. U.S. Natl. Herb. 37: 161-396. Rickerr, H. W. & CorrAsoRATOns. 1967. Wild Flowers of the United States. Vol. 2, The Southeastern States. McGraw-Hill, New York. [Onagraceae, pp. 304-315.] SATTLER, 1973. Organogenesis of flowers. Univ. Toronto Press, Toronto. [Myrtiflorae, . 112-119] SESHAVATARAM, V. 1970. Onagraceae, p. 220-225. In Proceedings of the Symposium on Comparative er A x Angiosperm held at Delhi from May 8 to 18, 1967. [Bull. Indian iu Sci. Acad. N SKVARLA, J. J., H. RAVEN, x r2 CuissoE & M. Smarr. 1977. An ultrastructural study of viscin T ads in Onagraceae pollen. Pollen & Spores (in press STEBBINS, C. L. 1967. Adaptive radiation and trends of evolution in ri higher plants. Evol. Bi iul. 1: 101-142. ————. 1974. Flowering Plants: Evolution Above the Species Level. Harvard Univ. Press, bridge. SuBRAMANYAM, K. 1951. Studies on the relationships of the Melastomaceae. Mysore Univ. Half-yearly J. N B, 12: 327-330. & L. AYANA. 1969. A contribution to the floral anatomy of some members * J. Jap. Bot. 44: TAKHTADZHYAN AK HTAJAN], A. 1966. Sitona i Filogeniya Tsvetkovykh Rastenii. Izda- teľstvo “Nauka,” Moskva. [Issued 1967.] 197 Evolution und Ausbreitung der Blütenpflanzen. G. i ag — t. VLIET, G. J. C. M. van. 1975. Wood anatomy of Crypteroniaceae sensu . J. Microscop. 104: & P. 7 1975. Comparative anatomy of the Crypteroniaceae sensu lato. Blumea 22. Ve 195. Wurpack, J. J. 1971. Certamen Melastomaceis XVII. Phytologia 21: 353-368. The previous issue of the ANNALS or THE Missourt BOTANICAL GARDEN, Vol. 64, No. 2, pp. 145-380, was published on 2 February 1978. 656 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Preliminary Announcement THIRTEENTH INTERNATIONAL BOTANICAL CONGRESS Sydney, Australia. 21-28th August, 1981 THE PROGRAMME will consist of 12 sections—molecular, metabolic, cellular and structural, developmental, environmental, community, genetic, systematic and evolutionary, fungal, aquatic, historical, and applied botany. There will be ple- nary sessions, symposia, and sessions for submitted contributions (papers and posters). Chairman of the Programme Committee: Dr. L. T. Evans. Frgrp Trips will include visits to arid and semi-arid regions, eucalypt forest, rain forest, heath, coastal vegetation (e.g. Great Barrier Reef, mangroves) etc., and specialist trips. Chairman of the Field Trips Committee: Prof. L. D. Pryor. First CIRCULAR, containing details, will be mailed in August, 1979. Send your name and full address, preferably on a postcard, to ensure your inclusion on the mailing list. Enquiries should be sent to the Executive Secretary, Dr. W. J. Cram. ConcREss AppRESs— 13th I.B.C., University of Sydney, N.S.W. 2006, Australia. Sponsored by the Australian Academy of Science. blic cation from is Amable in MICROFOR © COMPLETE | “INFORMATION | „ uw Publications the . ee PM GARDEN The Awars OF THE i BOTANICAL Soe contains « con- : tributions mainly in plant systematics. The ANNALS appears four times a year, and four numbers constitute a volume. Many back issues are available. Tue deci below a are siete net; j thare is no E dicount, to ne. 3 : tesa eet Dr mail to por. sys senate early volumes contain numerous papers on topics ranging from borde metabolism i in _ Azotobacter to pod and stem blight of soybean. Volumes 1-15 con- tain a series of 15 papers on North American Thelophoraceae by E i: E. A. Burt, and B. M. Duggar Published a an pond mers on He mf e. of viral particles in volume 8. The first 55 volumes (1914-1968) of de Asas are now iu d UE dexed. The INDEX catalogues the 769 titles contributed. by We authors in these yo nes. iss INDEXi is soft bound and costs bis 00. 5 oe F VvoluME 64 SUM | | 1977 NUMBER 4 MISSOURI E RI BOTANICAL L GARDEN - jua ER ANNALS contains papers, primarily in systematic boten = 3 x n from the Missouri Botan ical Garden. Papers EUH outside the Garde 3 — > editor for inf the ANNALS. — EDITORIAL COMMITTEE | Missouri Botanical Garden Published four times a year by the Waa Botanical Garden » Pres ss, St. Louis, Missouri 631 10. ae For Subscription information contact the Business Office F o Annals, x 368, RA 1 New 5 Lawrence, Kansas 6 66044. ANNALS MISSOURI E BOTANICAL GARDEN VOLUME 64 1977 NUMBER 4 PERSPECTIVES IN TROPICAL BOTANY: INTRODUCTION P. B. ToMLINSON! AND PETER H. RAVEN? The following papers were presented at the Symposium entitled “Perspec- tives in Tropical Botany,” held 22 August 1977 during the 28th Annual Meeting of the American Institute of Biological Sciences, Michigan State University, East Lansing. It was cosponsored by the Botanical Society of America, Ecological Society of America, American Society of Naturalists, and American Society of Plant Taxonomists. Intensified studies of the plants, vegetation, and ecosystems of the tropics is not some esoteric or arcane aspect of pure science but essentially an area of applied biology which is much neglected. A number of familiar factors contrib- ute to a feeling of urgency among tropical plant biologists which lead to this small symposium: (a). The greatest concentration of floristic and functional diversity occurs in the tropics. (b). There is a relative dearth of active research scientists in the field of tropical botany. (c). Tropical ecosystems are being destroyed at a rapid rate without ade- quate compensation in terms of conservation of representative vegeta- tion types and of genetic resources. The total effect is one of an overall deficiency in our understanding of bio- logical processes in plants, which is unfortunate since it occurs within countries which have predominantly agriculturally based economies with plants as a major natural resource. This message is stated to the point of tedium, but it needs constant attention by professional biologists. This symposium emphasized that this imbalance presents a problem of uni- versal concern. Tropical botany is not a discipline set apart from the rest of ! Harvard University, Harvard Forest, Petersham, Massachusetts 01366. ? Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. ANN. Missouni Bor. Garp. 64: 657-658. 1977. 658 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 plant science—it is an essential aspect of the whole subject, and for this reason any deficiency in our understanding of tropical vegetation is detrimental to our total understanding of the biology of plants. Any comparative study of plant form or structure is based on incomplete premises if it does not take into con- sideration the total range of natural variability in plants, i.e., including the enormous diversity of plant life in the tropics. Any physiological mechanism or ecological response has been analyzed insufficiently if it does not consider how these mechanisms or responses are mediated in tropical climates and especially in the nonseasonal climates of the lowland tropics. Any evolutionary idea has been incompletely scrutinized if it has not been tested against tropical examples. Any generalizations of plant population biology must apply to the frequently distinctive composition of tropical forests. And yet this cosmopolitan consideration is frequently lacking because of the enormous geographical paradox that besets biology—namely, it is temperate based, with its practitioners trained and deployed within relatively depauperate temperate ecosystems. The paradox is heightened by the fact that the greatest repository of known information about tropical plants resides in the herbaria and libraries of temperate countries. The object of the Symposium was to call upon a group of contributors, chosen because of their wide research experience in the tropics, and ask them to comment upon the present status of an aspect of tropical plant science wherein they are acknowledged experts. The further suggestion was made that they assess future needs in tropical research, although this was not a prime requirement. Tropical biology has developed enormously in the United States since World War II partly by the efforts of a number of institutions which have emphasized the discipline and particularly by the concerted action of a group of Universities that led to the formation of the Organization for Tropical Studies which has pro- vided a mechanism whereby a generation of biology students have been intro- duced to the tropics. An overview of an important aspect of tropical science is therefore appropriate at this stage, and we hope that the limited opportunity for presentation which this Symposium permits will generate further interest and action in the field of tropical botany. Whether we look back over what has been achieved or forward to what has still to be done, the opportunity for a brief appraisal is a welcome one. The twentieth century draws rapidly to its end—a century in which scien- tific advance has wrought endless miracles, culminating in the technological expertise which allows us to explore the universe directly. How unbalanced might our perception become before we discover that too little has been done too late to allow us to understand our more immediate environment, which after all provides us with food, energy, shelter, and an abundance of natural, renew- able resources—the environment provided by the plant kingdom. FLORISTIC INVENTORY OF THE TROPICS: WHERE DO WE STAND?! GHILLEAN T. PRANCE? In a review of the vast topic of the inventory situation in the entire tropics I can only skim over the surface. I have aimed to pinpoint a few of the signifi- cant contributions (many other important ones are omitted) and to draw atten- tion to some of the areas in need of further work. These include both geograph- ical areas that are poorly collected and disciplines which are still neglected in our basic survey of the fascinating vegetation of the tropics. There is still a great deal to be done and time is running out as the natural vegetation is being destroyed. Brazier et al. (1976) said: “More efficient use of the natural tropi- cal forest could be achieved if sufficient information on the extent, composition and structure of the resources were available The data which we collect from botanical inventory are not only useful for the study of floristics and evolution, but are also of vital importance for both conservation and utilization of the tropical vegetation. Or WHAT ARE WE MAKING AN INVENTORY? Table 1 gives a summary of the estimated number of species in the different major plant groups, compiled from the best available sources. The tropical flora consists of some 155,000 species of flowering plants, 11,000 ferns and fern allies, 16,000 bryophytes, and at least 90,000 fungi. The tropical flora is by far the richest in species diversity, yet it is also the most poorly collected. This diversity is being reduced before we have made an adequate basic inventory let alone conducted modern biosystematic and population biology studies in the area. Even to understand the origin and dynamics of our temperate flora, it is essen- tial to have adequate knowledge of the tropical flora from which the temperate flora was derived. Tropical Africa has the smallest number of angiosperm species, 30,000, in- cluding various islands and the 10,000 species of Madagascar (Koechlin, 1972). Tropical Asia, Australia and the Pacific have at least 35,000 species, and tropical America has about 90,000 species or 37.5% of the worldwide total. Unfortu- nately, the state of knowledge of these floras is also inversely proportional to the species diversity, with the American tropics much more poorly known than the African and Asian tropics. In any discussion of inventory of the tropical flora it is important to consider habitat diversity and species diversity. We tend to ignore the habitat diversity of the tropics which contributes to its species richness, and to think of it as one J am grateful to the many people who have helped to provide information about the areas of their specialty especially to Mr. F. N. Hepper, Drs. F. R. Fosberg, M. Jacobs, Alain er, W. Meijer and A. Gentry. I thank Mr. W. C. Steward for much bibliographic assis- bet Mrs. F. Maroncelli who typed the manuscript, and Drs. Howard Irwin and Scott Mori for reading the manuscript critically. The New York Botanical Garden, Bronx, New York 10458. ANN. Missouni Bor. Garp. 64: 659-684. 1977. 660 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE l. Estimated species numbers of major plant groups in the world and in tropics. Data compiled from many sources such as Ainsworth (1961: 405-407), Jacobs (1974, 1977), and Raven (1976a). Tropical Tropical Tropical Taxa Worldwide Tropical Africa sia _ America Flowering Plants 240,000 155,000 30,000 35,000 90,000 Fungi 120,000* 90,000 20,000 20,000 50,000 Ferns 12,000 11,000 1,000 6,000 5,000 Mosses 12,000 9,000 — — — ea 11,000 7,000 — — — 17,000 — — — — Lichens 16,500 — — — 2,500 20 ther estimates - for fungi are as as high as 250, 0,000 sp species, e. g., Martin (1951), Jones (1951), and Rogerson (pers. comm.). large uniform rain forest. In fact, the tropics contain many arid regions with deserts or scrubland, such as the caatingas of northeastern Brazil, a large tem- perate element in the flora of high mountains, and a unique alpine flora such as that of the páramos in South America and the Afro-alpine region described by Hedberg (1964), besides many different types of forest and savanna. The habitat types of Malesia were summed up by Jacobs (1974), those of South America by Hueck (1966), and those of Africa by White (in press) in a book to accompany the second edition of the UNESCO vegetation map of Africa. There is not time here to summarize the fascinating diversity of habitat in the tropics, but it is important to collect from and to plan conservation of this habi- tat diversity. Until now collecting has given rather uneven coverage to the different habitats. The location of different habitats has been overlooked fre- quently in biogeographic analyses of the neotropical vegetation, although the inventory of habitat distribution is vital to biogeographic studies. WHAT IS LEFT To INVENTORY? The tragedy of the biological inventory of the tropics is that destruction of the vegetation is proceeding more rapidly than the inventory. The tropical flora occurs mostly within the territory of developing countries where technological advance is urgent. Such advance traditionally includes the destruction of large areas of natural vegetation for replacement by farms, timber concessions, devel- oping towns, etc. In addition there is population pressure in many tropical coun- tries where the annual net population increase is often over 3% (see, for exam- ple, The Environmental Fund, 1976 Many authors have drawn attention to the destruction of the natural vegeta- tion in the tropics, for example, Gómez-Pompa et al. (1972), Richards (1973), Janzen (1974), Holdridge (1976), Myers (1976), Raven (1976b), Gentry (1978b), and many of the authors in Prance & Elias (1977). It is not the purpose of this paper to review in detail the destruction of the tropical vegetation, but as the tropical areas are vital for the understanding of the biology and evolution of all plants, it is important to draw attention to the urgent need to accelerate all biological inventory and conservation work in the tropics. According to recent — 1977] PRANCE—FLORISTIC INVENTORY 661 estimates 49.2 acres of tropical rain forest are being removed each minute or a total of 11,000,000 hectares a year (Lucas, 1977; Sommer, 1976). Inventory work daily becomes a more important task to perform, as destruction of natural habi- tats encroaches. Since there is not a separate treatment of conservation in this symposium, I feel that it must logically be stressed as part of the inventory. It is not possible or profitable to list examples of tropical destruction from each area discussed below, but I draw attention to this race between inventory and destruction in the tropics and hope that we can also focus our efforts more towards conservation. None of the other subjects treated at this symposium can be completed without the conservation of large areas, and without a compre- hensive basic inventory. Tue REGIONAL STATUS OF INVENTORY AFRICA Progress on the status of systematic work in tropical Africa is readily acces- sible through the publications and symposia of the “Association pour l'étude taxonomique de la flore de l'Afrique Tropicale” (AETFAT). This organization publishes an annual index which includes a bibliography and lists of new taxa and nomenclatural changes for all tropical African plants. Progress reports on collections, the regional floras, mapping, etc., are given in the proceedings of their symposia which take place every fourth year (see, for example, Hedberg & Hedberg, 1968; Kubitzki, 1971). AETFAT plays a similar role for Africa as Flora Malesiana does for Asia in making available much information and biblio- graphic data invaluable for research in the area. A review of the current status of collecting in tropical Africa was given by Hepper (in press). Léonard (1975) prepared, for AETFAT, a map of the extent of floristic exploration in Africa south of the Sahara up to 1963. This map divided the region into 3 categories: poorly known, moderately known and well-known areas. Hepper (in press) gave up-to-date information of changes to this map and a revised edition will be presented at the 1978 AETFAT Congress. The Flora of Tropical Africa (Oliver, 1868-1937) is the only attempt at a general flora of the region. This has been largely replaced by the modern regional floras, especially the Flora of West Tropical Africa (recently revised), Flora of Tropical East Africa, and Flora Zambesiaca. Current African floras are summarized in Table 2. Statistics for the description of new taxa in Africa from 1953-1965 were summarized by Léonard (1968) and showed a gradual decline from 1,177 new names (577 new species) in 1953 to 723 new names (287 new species) in 1965. The rate of description of new species continued at approximately the same rate in 1971-1975 and is shown in Fig. 1. The fact that 270 new species were de- scribed in 1975 shows that the basic species inventory of the African flora has not yet ended. Figure 1 also shows the amount of synonymy proposed in the years 1971-1975 (data from the AETFAT indices). It shows that there is appar- ently a gradual drop in the net gain in species because of increasing synonymy, 218 net gain in 1971 as compared with 119 in 1975. Nevertheless, the total 662 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE 2. Principal regional floras of tropical Africa. Publication Editor or Author Flora of West Tropical Africa ed. 1 TER & Dalziel ( 1927- 1936) ed. 2 revised by Keay et al. (1954-1972) Flora of Tropical East Africa Turr vil & Milne- Redhead (1952-) Flora Zambesiaca Exell & Wild (1960- ) Flora of Egypt Laurent-Tackholm | Ira Flore du Sénégal Berhaut (1954 37) Flore du Gabon Aubréville (1961-) Flore du Cameroun Aubréville (1963-) Flore du Congo, du Rwanda et du Burundi Robyns (1948-) Syllabus de la Flore du d van da Troupin (1971) Conspectus Florae Angolen Carrisso ( 1937— Prodromus einer Flora von "Südwestafrika Merxmiiller ( 1966- ) Flora of Southern Africa Codd et al. (1963-) Flore de Madagascar et des Comores Humbert (1936-) number of new descriptions in Africa south of the Sahara for the 21 year period, 53-1973, are impressive: 391 new genera, 7,478 new species, and 2,538 infra- specific taxa. That is a new genus every 3 weeks and a new species for every day of the 21 years (data from Hepper, in press). A flora in this active state of description that is still adding 1,000 new species over a five-year period is obvi- ously also in need of further collecting. Many of the reports on the progress of various African floras given in Kubitzki (1971) include emphasis on the need for further collecting, for example, Boulos (1971) for Libya, Aké Assi (1971) for the Ivory Coast, Le Thomas (1971) for Gabon, etc. Distribution maps of African taxa such as those published by Bamps (1969) in the very useful series "Distributiones Plantarum Africanarum" (see Fig. 2), show that the African flora is really well explored in comparison with the Neo- tropics. For this reason more analytical phytogeographic papers have come from taxonomists working on the African flora. The better known plant distributions have enabled much better phytogeographic analysis of the flora, see, for exam- ple, White (1962, 1965, 1971), the introductory chapters in Chapman & White (1970 Hepper (in press) summarized the collecting situation in Africa as still hav- ing large gaps. He said that general collections are now required only from lesser known regions, and he stressed the need for specialist collections and for resident botanists to carry out long-term investigations. He pointed out some particular gaps in collecting such as the tendency to collect mountain tops and ignore the forested slopes. For further information about Africa the reader is referred to Hepper's paper. The native African flora has been disappearing rapidly under human popu- 5 Ficure 1. The 3 of new species in tropical Africa: data from the AETFAT Index, 1971-1975. The open area represents the total number of new species described, the cross hatched area CIE the number of species names reduced to synonymy. The graph in the center represents the net gain in species each year. IN o wn — 664 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Distributiones planturum africanarum, 10 (31-12-76) — | , 2: : . | PED SEN A " ua | I 4 ^ f Chrysobalanaceae 327. - Parinari Aubl. (1775) F. White Synthesis tabularum 328-334 Distribution of the genus Parinari Aubl. ( Chrysobalanaceae) in Africa: From White [t2] F Plantarum Africanum 10: 281-334. The dots represent presence a degree 1977] PRANCE—FLORISTIC INVENTORY 665 lation pressure for longer than that of Malesia or the neotropics. Shantz (1948) discussed the shrinkage of the tropical forests of Africa and Shantz & Turners (1958) photographic account of the destruction of the vegetation of Africa is a frightening report for any biologist. Hepper (in press) also stressed the urgency for collecting the poorly known areas because of the rate at which the natural vegetation is disappearing. In Madagascar, where the largest contributions have been made by H. Perrier de la Bathie and H. Humbert, the original species-rich forests have been almost totally disturbed (Gentry, pers. comm.), and the race to collect this exciting flora before it is obliterated is lost. Koechlin (1972) summarized the situation in Madagascar: “Many problems still have to be solved in the field: although the exploration of the flora is well advanced, much remains to be done in the areas of plant biology and phytosociology.” TROPICAL ASIA The Asian tropics are probably not as well collected as Africa, but are much better known than the neotropics. Much information about tropical Asian bot- any has been compiled and published in Flora Malesiana and its extremely use- ful Bulletin, largely through the initial efforts of Professor C. G. J. van Steenis. The Flora Malesiana project covers Indonesia, Malaysia, Brunei, the Philippines, Singapore, eastern New Guinea, and the Solomon Islands. An excellent sum- mary of the botanical status of the region has been given by Jacobs (1974). The situation has changed little since that report. The history of collecting in Malesia is given by Mrs. van Steenis-Kruseman (1950) in a "Cyclopaedia of Collectors" published in volume 1 of Flora Malesiana and updated from time to time in the Flora Malesiana and in the Bulletin, for example, van Steenis-Kruseman (1958, 1974, 1977). It is not, therefore, necessary to repeat the data given in these sources but rather to indicate some of the gaps in collecting as given below. Dr. Jacobs (pers. comm.) lists the following places in Malesia as undercol- lected and in need of further basic inventory: The Andaman Islands, Southern Sumatra, Central Borneo, Celebes, Kabaena, Ceram (expedition planned), West New Guinea (especially Meerilakte and the Star Mountains in West Irian), the Kikari area in the south of Papua, the Philippines (especially the Sierra Madre on the east coast of Luzon), and the Cape York peninsula of Australia which has in recent times yielded several genera that were known only from Malesia. The northwest Australian coast is still poorly known. Perhaps Celebes is the least collected area and is now less known than New Guinea, especially the eastern and southeastern area of the island. Celebes also illustrates the race against de- velopment, since the International Nickel Company in cooperation with Bechtel has a billion dollar nickel mining concern in Celebes. Table 3 from Flora Male- siana reproduces their synthesis of the collecting situation in the larger units of Malesia. It points to the need for further collections from Sumatra and Celebes. Jacobs (1977) has summarized the progress in the publication of Flora Male- siana. By the end of 1976, 116 families, 453 genera, and 3,288 species of angio- sperms had been monographed out of an estimated total of 25,000 species. The fact that only 13.15% of the flora has thus far been published, together with the 666 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 8(1): 3 TABLE 3. Collecting density (specimens/km*) of Malesia (from Flora Malesiana, Ser. 1, . 1974.). Density Density Surface Pes 8 Index | (km?) to 19 1972 1950 — 1972 1. Sum 479,513 87.900 99 .000 TM 21 9 Malay Pe ninsula 132,604 191,055 232,000 145 175 3. Java 32,474 247,522 260,500 187 197 4. Lesser Sunda Is. 98,625 24.545 36,000 25 36 5 )rneo 739,175 91,550 194,200 12 26 6. Philippine Is. 290,235 180,090 200,000 62 69 7. Celebes 182,870 32,530 ; 8 19 8. Moluccas 63,575 27,525 30,400 43 48 9. New Guinea 894,855 106,775 233,000 12 26 Totals 3,013,926 989,492 1,319,100 += 83 £ = 44 figures for Flora Neotropica given below, shows the magnitude of the task in tropical areas and the shortage of botanists to work up the results of inventory. The slow production of monographs is a serious problem and lags behind the progress of development. However, a basic collecting inventory is more impor- tant before forests are destroyed. For Pteridophytes, Flora Malesiana has pub- lished 5 families, 14 genera, and 350 species or 14% of the estimated 2,500 species. A comparison of the three major continental areas of the tropics in terms of statistics of species descriptions is not as straightforward as it may seem since the different status of knowledge in each flora has tended to result in a rather different species concept in each area. Although the tendency is toward much new synonymy in all three areas, the Malesian botanists seem to have a more conservative attitude to the species concept. For example, Leenhouts (1967) reduced all 255 species of Allophylus (Sapindaceae) to the single species A. cobbe (L.) Raeusch. Whitmore (1976) cites other examples. The concept of the reticulately polymorphic ochlospecies came from work on the African flora (White, 1962). The species concept in Africa lies somewhere between that of Malesia and the narrower concept that has predominated in the neotropics until recently. It is not the purpose of this paper to evaluate the merits of these dif- ferent concepts, but an acknowledgement of their existence is necessary for a comparison of data between the different areas. Whitmore (1976) also pointed out the different kinds of species that exist in the tropics, accepting three kinds; the discrete, isolated and morphologically invariable species, the species with distinct infraspecific taxa, and the reticulately variable ochlospecies. There are numerous local floristic works within the Flora Malesiana region, the best known of which is Backer & Bakhuizen van den Brink’s (1963-1968) Flora of Java. In the Asian tropics outside Malesia the situation is similar with a reasonable basic inventory but still some neglected areas. India and Burma have had much less collecting since World War II, but a botanical survey of India is making good progress. In India there is a general reluctance to collect trees in primary vegetation and little specialist collecting 1977] PRANCE—FLORISTIC INVENTORY 667 TABLE 4. Collecting status of some Pacific Islands: (1) only casual collecting; (2) oorly collected (not professionally collected); (3) moderately well collected; (4) rather well collected but some gaps; (5) quite wel collected. More than one 5 means differ- ent islands in groups collected to different degrees. (Data from F. R. Fosberg, pers. comm.) Collecting Island Status Revillagige de 3 Cook Islands 2,3,4 Coco 2 Northern Cook Islands 3,4 cM am 4 Wake Island 5 Easter Island 4 Marshall Islands, Northern 4,5 Hawaiian Islands 4 Marshall Islands, Southern 3,4 Phoenix Islands 5 Gilbert Islands 2,3,4 Other = i Pacific atolls 4 Naura and Banaba Marques 3 Ellice Isl 2.9 Society a (high ) 2,3 Niue Elend 4 Society atolls 2, 3, 5 Rotuma Island 4 d Islands 2 3,4 Wallis and Horne Islands 2 Makatea 2 Tokelan Island 3 Awana Islands 3,4 Samoa Islands 4 Ray 3 Tonga Islands 2,3,4 has been done. Some local floras are replacing Hookers (1872-1897) Flora of India, as for example the recent Flora of the Hassan District by Saldanha & Nicolson (1976). Burma has had the smallest portion of its flora collected. Thailand has had intensified general collecting since 1960 with a wide cov- erage of habitats and areas but little specialist collection. Collection has been stimulated by the joint Thai-Danish project on the Flora of Thailand under the leadership of Kai Larsen and Tem Smitinand and their collaborators. Sri Lanka has been extremely well collected and worked up under the Flora of Ceylon project directed by F. Raymond Fosberg. This project has included much specialist collecting and the results of this are obvious in the resultant monographs. ndo-China has had little collection since World War II except for a few vegetational studies in Laos and South Viet Nam. The political upheaval in that region has not been conducive to botanical inventory. The use of chemical de- foliants in the war has truly devastated large areas of the forests of Indo-China. The Flora Malesiana Bulletin serves a very fine role of reporting on progress in tropical Asian botany, even in countries outside the range of the Flora itself, and is commendable for the amount of useful information generated. The annual columns on progress in Malesian botany, expeditions and exploration, and on recent publications serve to keep us up to date on the state of Asian botany. There is a need for such a bulletin attached to Flora Neotropica. For the tropical Pacific islands Dr. F. R. Fosberg has provided me with the data presented in Table 4. It shows that there are islands which remain poorly collected. Exact statistics on collecting are not available, but there is obviously much still to be done in this area that is so fascinating from the point of view of island biogeography. Dr. A. C. Smith has worked extensively in Fiji so that the archipelago can now be considered well collected, and he is following up with a flora of the islands. 668 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 THE NEOTROPICS The New World tropics are certainly much less known than Africa or Asia and are still in the process of the first basic inventory. New species are still being collected in large numbers from many places, as, for example, the large number of new species from recent collections in the uplands of Panama, from coastal Ecuador, from the forests of the state of Bahia, Brazil, and from many other localities. The collecting density throughout the neotropics is much less than for Ma- lesia, but added to this, the greater number of species in the neotropical flora and the unevenness of collection throughout the area mean that the basic com- prehensive species inventory is still most inadequate and by no means nearing its completion. Unfortunately there is no equivalent of the Flora Malesiana Bulletin or the AETFAT publications in the neotropics. Thus, calculations of botanical activ- ity are harder to make and are less accurate. We hope, however, that the Orga- nization for Flora Neotropica will gradually begin to fill in this information gap as it begins to diversify its interest from only producing taxonomc monographs. A comparison of many aspects of the ecosystems of Africa and South Amer- ica is given in Meggers et al. (1973), but it does not cover the subject of inven- tory in any detail. The last comprehensive review of the state of neotropical botany was that of Verdhoorn (1945). Much collecting has taken place since 1945 and some aspects, particularly from the conservation point of view, were surveyed i Prance & Elias (1977). In Prance (in press) I give a country-by-country review of the status of botanical exploration in South America, and Gentry (19782) reviewed the floristic needs of Central America and the Pacific coastal region of northern South America. There is no space to give such a detailed review here, but a few examples will serve to show the situation in the neotropics. Figure 2 is a distribution map of the pantropical genus Parinari in Africa from White (1976). The dots on the map represent presence in a degree square. Figure 3 is a similar distribution map of the same genus in the neotropics. The genus is common and widespread on both continents. It can be seen how much more densely the map of Africa is covered. Judging by the frequency of the individuals I have encountered in fieldwork in the neotropics and by the num- ber of habitats occupied by the different species, I would predict that to be accurate the neotropical map should be almost as densely covered with dots for each degree square as is Africa, and that the distribution difference is actually the result of inadequate collections in South America. Gentry (1978b) reported on an expedition to Cerro Tacarcuna, a previously unexplored mountain on the Panama-Colombia border. At least 20% of the 239 species collected above 1,400 m have turned out to be new. A similar figure is E URE 3. Distribution of the genus Parinari Aubl. (Chrysobalanaceae) in South Amer- The dots = presence in a degree square similar to the African distribution map pede used in Fig. PRANCE—FLORISTIC INVENTORY 669 1977] 400 — 300 — 400 500 FOO MULES iL SINUSOIDAL PROJECTION WEST LONGITUDE SOUTH AMERICA CHRYSOBALANACEAE . (1775) 18. - Parinari Aubl Synthesis of plates 1-17 670 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ABLE 5. New species and subspecies of Chrysobalanaceae described since the mono- graph of the family in 1972 ( Prance, 1972a) Date of collection € Species Locality of the t N dolichopoda Disa Data 1972 Couepia 8 1971 Couepia glabra Brazil—Amazonas 1971 idein marlenei Brazil—Amazonas 1972 Hirtella arenosa Brazil—Amazonas 1968 Hirtella conduplicata Brazil—Amazonas 1973 Licania aracaensis Brazil—Amazonas 1975 Licania chiriquiensis Panama 1975 Licania furfuracea Venezuela—Bolivar 1975 Licania jefensis deben 1969 Licania jimenezii 9 1 1971 Licania marlenei Brazil —Anizons 1972 Licania montana Venezuela—Lara 1975 Licania morii Panama 1975 Licania octandra (Hoffmgg. ex R. & S.) Brazil—Amazonas 1973 Kuntz pn grandifolia Licania 5 nsis Venezuela—Bolivar 1973 Licania 1 Brazil Amazonas 1974 Licunia „ Peru Loreto 1974 Licania sp. 1 Ecuador 1969 Licania sp. nov. 2 Panam 1975 Licania sp. nov. 3 razil—Amazonas 1976 ‘ania sp. nov. 4 Colombia Valle 1972 Licania sp. nov. 5 Panama 1972 Hirtella sp. nov. 1 Peru—Lore 1976 Hirtella sp. nov. 2 Br ZI os 1976 true of a recent collection of J. Murça Pires in Amazonas, Brazil, from the re- cently discovered and isolated sandstone peak Serra Acará. The number of new species that are still being described from recent collections is indicative of the state of collecting. In 1972 I monographed the neotropical Chrysobalanaceae (Prance, 1972a). The monograph recognized 328 species in the 8 genera. Table 5 lists the 26 new species that I have described (or are ready to be described ) in the five years since I completed the monograph. The new species amount to 7.93% of the original number of species. All except three of the new species are based on type collections made since 1970. Many other monographic, floristic, and descriptive works on the neotropical flora, especially from Panama south- ward, are adding species at a similar rate. The number of new species in Table 9 from the forests of Panama and from Amazonia points to two of the under- collected areas of the neotropics, although there are also many other places out- side the main distribution range of the Chrysobalanaceae. In addition to these new species, large range extensions of several of the original 328 species have occurred. For example, Licania affinis Fritsch was reported as a species con- fined to the Guianas. It has recently been collected several times in Panama, adding another species to the increasing list of Guiana-Panama disjunctions reported in Gentry (1975). While this is a true disjunction, many other species previously thought to be of local distribution are now seen to have much wider 1977] PRANCE—FLORISTIC INVENTORY 671 continuous distributions. For example, in Prance (1972a) I cited Couepia lon- gipendula Pilger as endemic to the Manaus region of Amazonia. In 1973 I collected it over 1,200 km away on the Rio Curicuriari. It was then collected later in 1973 on the Rio Cunhua also over 1,000 km from Manaus to the south- west rather than northwest, showing that this species is actually quite wide- spread in Amazonia (Fig. Gentry (1978b) 1 a similar case in a species of Siparuna known only from Panama and western Ecuador, 1,500 km apart, but subsequently collected in Chocó in Colombia. The number of species added to the Chrysobalanaceae and other groups in recent neotropical monographs is in marked contrast to the situation in African and Asian Chrysobalanaceae where few new species are being discovered. Ja- cobs (1974) cites a good example from the Malesian families monographed by Dr. P. W. Leenhouts (see also Leenhouts, 1976). The following additions to Flora Malesiana monographs have been made: Burseraceae (108 species), additions after 14 years: 1; Connaraceae (38 species), additions after 12 years: 0; Dichapetalaceae (15 species), additions after 13 years: 0; Goodeniaceae (8 species), additions after 13 years: 3. The neotropics, as reflected in Flora Neotropica has a much less complete species inventory. In 1972 I monographed neotropical Dichapetalaceae (41 species; Prance, 1972b), since that date I have described 3 new species showing the same trend as for neotropical Chrysobalanaceae. However, this comparison of species between Malesia and the neotropics is further complicated by the differences in species concepts referred to earlier. Although many new species are being added to the neotropical flora, also much synonymy is taking place. For example, in the monograph of Chrysobala- naceae (Prance, 1972a), where 75 new species were described, 76 names were placed in synonymy, giving an almost even result. Monographic treatments are finding many “regional” species. I suspect that in Lecythidaceae, which I am currently monographing with Dr. Scott Mori, the percentage of synonymy will be even higher since the last monograph by Reinhardt Knuth (1939). Knuth was a renowned splitter, and did not rectify the species described from several different regions under different names. The dangers of such an incomplete inventory are obvious. Extreme caution must be taken with drawing biogeographic conclusions from plant distributions as we know them today. Some disjunctions are now well established and have logical historic bases, such as the climatic changes in the Pleistocene and post- Pleistocene ( Haffer, 1969; Prance, 1973; Toledo, 1976); others are artifacts of a poor collection sample. Such disjunction as Panama-Guiana or Amazonia to the coastal forests of Bahia are well established. Lamentably, the destruction of the forest is proceeding at such a rate that we may never be able to put together the accurate distribution pattern of many neotropical plants. Central America is one of the better known regions of the neotropics with 672 ANNALS OF THE MBSOURI BOTANICAL GARDEN [Vor. 64 SOUTH AMERICA SINUSOIDAL PROJECTION 40 30 WEST LONGITUDE 20 | Ficure 4. The distribution and range extension pattern of Couepia longipendula Pilger. 1977] PRANCE—FLORISTIC INVENTORY 673 TABLE 6. Collecting density in Central America, data from Gentry (1978a) and other sources. No. of Surface Area Density Index Country No. of Species Collections (km?) m' Guatemala 8,000 80,000 108,880 0.73 Belize 3,000 25,000 22,272 1.12 El Salvador 2,500 8,000 21,392 0.37 Honduras 5.000 40,000 112,079 0.36 Nicaragua 5,000 5,000 129,990 0.038 Costa Rica 8,000 70,000 50,695 1.38 Panama 8,000 100,000 75,643 1.32 regional floras existing or in preparation in most countries. Work on the regional floras such as Guatemala, Costa Rica, and Panama has stimulated much botani- cal collection. Table 6 gives the approximate collecting density for the region. Only in two countries does it reach as high as 1.3 specimens per square kilometer. This may be compared to Table 3 where the lowest density for Malesia is Cele- bes with 19 specimens per square kilometer. Such a table for South America would be even lower than that of Central America. Holdridge (1976) discussed the reasons for the diversity of the Central American vegetation. Perhaps the richest rain forest in the neotropics is that of Chocó, Colombia where annual rainfall is up to 10,000 mm (Lellinger & de la Sota, 1972). Only about 8,000 collections have been made from this area, first by Cuatrecasas and more recently by Gentry and Forero. It is certainly one of the most important and most interesting areas in the tropics for future collection. Colombia as a whole has the richest flora in South America with perhaps half the species of the neotropical flora occurring within her territory, 45,000 species (50,000 accord- ing to Schultes, 1951). Colombia as a country is still poorly collected and many areas rich in endemism such as Sierra de la Macarena and Sierra Nevada de Santa Marta are poorly known botanically. Venezuela is relatively well explored botanically in comparison to many other countries of South America, yet Steyermark (1974) stated that "less than 2% of Venezuela has been explored botanically." Maguire (1970) estimated that more than 75% of the flora of the Guyana highlands is endemic. With the high endemism from mountain top to mountain top in that region, even the intensive expeditions of Maguire, Steyermark, and other collaborators has just begun to inventory the flora of the sandstone mountain tops. The large state of Apure in Venezuela offers diverse habitats, savannas, lakes, rivers and rain forests, rarely visited by botanists. In the Guianas, French Guiana remains the least explored and is in need of much more intensive fieldwork. Svenson (1945) stated that "Ecuador is botanically one of the least known, though one of the richest countries in South America." This is still true today, although the Swedish-based Flora of Ecuador project has stimulated much more collecting activity in recent years. This has concentrated on the highland areas, leaving the Amazonian part of Ecuador still very poorly known. 674 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Peru also offers a wide range of plant habitats, from arid desert regions to the humid tropical forest of her Amazonian territory. Despite the long history of collection in Peru, it is still poorly known botanically. The Flora of Peru initi- ated by Macbride in 1936 has recently been reactivated as a cooperative Field Museum-Missouri Botanical Garden project. This has stimulated more collec- tions in recent years, particularly from the poorly collected Amazonian region. Brazil, the largest country in South America, has a long history of botany, and a great diversity of vegetation. With an area of 8,511,965 kme, the collect- ing density of the country is certainly well under 1 specimen per square kilo- meter. National herbaria have about 2 million specimens. Some of the poorly collected areas of Brazil include the state of Acre, Serra Pacaás Novas in Ron- dónia, the forests of northern Mato Grosso and Serra Cachimbo in Amazonia. Besides the Amazonian region there are many other neglected areas of Brazil such as the coastal forest of Bahia and Espirito Santo and some parts of the arid caatinga region. In January 1976 Brazil initiated an ambitious program called Programa Flora. This program plans to make a detailed inventory of Brazils vegetation by col- lecting programs and by the preparation of a computerized label data bank of Brazilian herbaria. The program is divided into five regional projects and Projeto Flora Amazónica has already begun. Arrangements for North American participation in the collecting program have been made and collecting will start in the fall of 1977. Bolivia is probably the least collected of all South American countries. Little collecting has been carried out since the time of the summary of collections from Bolivia by Herzog (1923: 14). The lack of a strong national botanical work in Bolivia has also hampered fieldwork by foreigners. There is a great need for collections from all over Bolivia. Paraguay, which lies geographically half within the tropics, is another poorly collected country where only about 30 collectors have worked extensively. Ar- gentinian botanists have visited Paraguay and made important collections there. There is very little primary vegetation left in Paraguay. The Caribbean islands of the Antilles have a flora of 12,000-15,000 species (Howard, 1977). The islands, which stretch over 1,700 miles east to west and 1,200 miles north to south, have many local endemics. For example León and Alain estimated that almost 50% of the 6,000 species of Cuba are endemic, and Hispaniola has 33% endemism in its flora of 5,000 species (Alain, pers. comm.). The history of floristic work has been a one-island approach which has led to many species being described from several islands, and more island “endemics” are being reduced than new species described. One of the needs of Caribbean botany is a monographic approach to compare elements of its flora with South and Central America and to calculate the true percentage of endemism. Howard (1977) has noted that "plant life of the Caribbean Islands cannot be regarded as unknown or needing immediate study or a massive collecting program." The area has been well collected in comparison to Latin America. There is, however, a need for any of the experimental type collections listed in the next section. Howard (1977) lists many examples of environmental de- 1977] PRANCE—FLORISTIC INVENTORY 675 TABLE 7. The most recent country or regional floras of Latin America. Country Status Reference Guatemala (almost complete ) Standley & "— (1958-) Belize Annotated checklist Standley & Record ( ) El Salvador Annotated checkl Standley & Calderón (1925) Honduras Various regional flor Standley (1930, 1931) ras and checklists shed Nicaragua Initiating di none publis Costa Rica Flora in progress Burger (1971 Panama Flora tad. completion Woodson & "w Na (1943-) Flora of Canal Z Standley (1928) Colombia Generic Flora eti none published Flora of the State of Pinto-Escobar (1966-) Cundinamarca in ke pangran Venezuela Flora in progre Lasser (1968-) Guyana No flor: Surinam Flora under revision Pulle (1932-) French Guiana ale ad a Flora, no modern Flora Lemée (1953) Ecuador Flora in Pus (6 families published) Harling & Sparre (1973-) eru Flora reactivated, in progress Macbride (1936-) Brazil No modern Flora since Martius s. Í Vianna (1965-) Various local floras e.g. Santa Catarina, Reitz (1965-) Res sag ai etc Hoehne (1940-) Bolivia No Flor Paraguay No Flora struction in the Caribbean, and, as elsewhere, further collecting data is needed for conservation information. Floras exist for many of the larger islands, for example, Cuba (León & Alain, 1946-1969), Puerto Rico (Britton & Wilson, 1923-1930), Jamaica ( Adams, 1972; Fawcett & Rendle, 1910-1936), and that of Hispaniola is in preparation by Alain. In spite of the better collecting status of the region, there are still novelties being found in the Caribbean, for example, Alain recently found a new species of the previously South American genus Talisia in Santo Domingo ( Alain, pers. comm.). Some of the smaller islands of the lesser Antilles have been overlooked. able 7 gives a list of the most important local floras in the neotropics. The collecting for the floras has done much to stimulate the inventory of the region, but most of the floras are based on inadequate specimen samples and conse- quently new species and extension of ranges must be added to the floras of each country. For example, the earlier family treatments of the Flora of Panama (Woodson & Schery 1943) are very inadequate in their species coverage and nomenclature. The recent exploration of the moist forests of Panama has added many elements to the flora. Another result of such an emphasis on regional floras has been the description of many species several times in the various re- gional floras. Even Paul Standley (1928), known as a "splitter," commented that “in tropical America . . . the flora has been studied from isolated centers with little regard for the species accepted at other centers, but with the assumption that each area is floristically distinct. Correlation through monographic work, covering a group throughout its range, will reduce the species that have been multiplied unnecessarily." 676 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE 8. Number of species described in Flora Neotropica monographs. Volume Author i mM _ Group | No. of Species hs iios 'ering Plants 1 Cowan (1968) Swart 127 2 Cuatrecasas (1970) 1 50 7 Berg (1972) Oln erasa en 68 8 Maas (1972) Costoideae 41 9 Prance (1972a) 5 328 10 Prance (1972b) Dichapetalacea 39 11 ance (1972c) dla 3 12 Prance & Silva (1973) Caryoc: 23 13 Rogers & Appan (1973) Manihot/ Manihotoides 99 14a Smith & Downs (1974 Pitcairnioideae 731 14b Smith & Downs (1977) Tillandsioideae 815 15 Morley (1976) Memecyleae 81 18 Maas (1977) Zingiberoideae 61 TOTAL 2,466 B. Fung 3 Singer (1970a) Omphalinae 52 4 Singer (1970b) Phaeocollybia 4 5 Singer (1970c) Strobilomycetaceae 13 6 Lowy (1971 Tremellales 148 16 Farr (1976) Myxomycetes 280 17 Singer (1976) Marasmieae 322 Tora. 819 Recent monographic work has shown the words of Standley to be true, and most neotropical monographs include a considerable amount of synonymy, but at the same time also include a large number of new species. Another stimulus to collecting in the neotropics is the Flora Neotropica monograph series initiated in 1964. Table 8 gives a list of the monographs pub- lished to date: 2,466 species or 2.74% of the estimated total of 90,000 flowering plants have been treated, and 819 species or 1.64% of the 50,000 fungi have been treated. Since the series also includes ferns, bryophytes, and algae, the task to be completed is enormous. Already new collections are outdating the existing treatments, see, for example, Table 5, the Chrysobalanaceae added since 1972. Maas (1977) contains a supplement to Maas (1972) which adds many new data. THE FUTURE INVENTORY In summary, Africa is the best collected continent of the tropics and is closely followed by Asia and Malesia. In these areas a basic inventory including most species exists, but the sample size of many species is still inadequate for a true understanding of their biology and ecology. In the neotropics the basic inven- tory is still underway, and many new species are still being found. There are many areas of South America still to be explored botanically. However, collect- ing should not now be slowed down anywhere in the tropics. A different empha- sis is needed now to provide an adequate experimental sample. Some of the foci 1977] PRANCE—FLORISTIC INVENTORY 677 for future collecting are outlined below, and these correspond with the needs of a more experimental approach to tropical taxonomy. Although herbarium inventory is still taking place in many areas, the experimental methods can often be carried out at the same time. For example, it is easy for any collector to carry fixatives and collect bud material for the study of chromosome numbers. Throughout the tropics many species are known from incomplete material. Future collecting should focus on previously inadequately collected material such as the fruits of many tall forest trees, and collections should be accompa- nied by good field data and notes on dispersal where possible. Jacobs (1976) pointed out that lianas are poorly collected and gave a succinct summary of collecting problems in lianas. The large fleshy, monocotyledons such as Zingi- beraceae (see Burtt, 1976), Musaceae, and Araceae are poorly collected, and pickled flowers are essential for adequate study. Much more liquid preserved material should be collected and distributed to specialists. At the outset of my work on Lecythidaceae it was necessary to obtain a large collection of preserved flowers over a period of several years before the complicated androecium struc- ture of the large fleshy flowers could be interpreted. Other groups that are poorly collected include tropical macrophytes, bamboos, palms, and Utricularia (see Taylor, 1977, for collecting techniques in Utricularia). Van Steenis (1977) made an important plea to tropical collectors to improve their field data. He pointed to the need for further label data on color, scent, size, texture, structure, and habit of specimens, for liquid material, and for black-and-white photographs of habit and habitat. As studies on floral biology, phytogeography, and evolution in the tropics increase these are needed. It is often better to collect fewer numbers but to document them well. Economic plants have often been neglected by taxonomist collectors who have the habit of leaving such things to economic botanists, agriculturists, or forest- ers. This has resulted in inadequate sampling of many of the most important economic plants and even their wild relatives. The contrary has occurred in some tropical areas where the local herbarium is a Forestry Herbarium. Col- lecting was concentrated on forest trees and “noneconomic” plants like forest herbs and lianas have been neglected, for example, in North Borneo, Surinam, and French Guiana where forest herbaria are the most active botanical institutes. Plants of secondary vegetation have always been neglected as “inferior cous- ins” of the primary forest. Some secondary areas have an extremely rich and interesting flora, and they should also be further collected. For example, many of the hard-to-collect forest lianas in Bignoniaceae, Malpighiaceae and Meni- spermaceae occur abundantly in the secondary forest areas of the neotropics. A survey by Rodrigues (unpublished data) found 374 species in 63 families on an area of 3,500 m? of secondary forest near Manaus. Various authors have drawn attention to the importance of secondary forest in conservation of primary areas. Thus, use of secondary areas for plantations can often relieve pressure on primary areas (Budowski, 1977). Secondary forest also played an important role in the evolution of the tropical flora ( Gómez- Pompa, 1972). It is, thercfore, most important that we make a better inventory of secondary areas in the tropics. 678 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 General collecting is important and has really provided the basic inventory of the tropical vegetation. However, a specialist in any family finds far more interesting things about his group than the general collector. The specialist soon learns to recognize his group from the diversity of the forest, and field studies by specialists have contributed many of the interesting results from the tropical forest. I have been accompanied by many specialists on my botanical expedi- tions and have often been impressed at their ability to find their groups, and the representation of any family in our collections always increases when there is a specialist present. There is much in favor of taxonomic focusing in collection. A general collector who concentrates on certain groups will also produce inter- esting collections. Another important aspect for tropical forest areas is the concentration on a small area over an extended period. This is best done by resident botanists and can be highly rewarding, from both a taxonomic and ecological point of view. For example the selection of one hectare of forest for study in a relatively well- known area near Manaus, Brazil yielded many interesting results, including at least two new species from the 236 tree species on the hectare (Prance et al., 1976). The detailed botanical study of Barro Colorado Island in Panama ( Croat, in press), was based on much fieldwork and treats 1,400 species from an area of 14.8 kme. This study has also enhanced many other interdisciplinary studies and is a good example of the usefulness and importance of minifloras and treatments and inventories of small areas of the tropical forest. An area where the individ- ual trees have been identified soon becomes the focus of many other studies apart from the original botanical inventory. Often the biggest problem facing other tropical biologists is the lack of such well-inventoried areas for their re- search. When we had inventoried the hectare of forest near Manaus, we were soon followed by entomologists, soil zoologists, and mycorrhiza specialists who could link their work to an accurate botanical inventory. Too many detailed tropical forest inventories in the past were carried out by foresters who relied on local names and did not collect enough herbarium specimens to document their inventories. There is a need for further well-documented inventories of small areas from throughout tropical forests. This type of inventory is quite as important as general collecting and often yields data of great use for conserva- tion, as well as ecology and other disciplines. It is one of the best ways to en- courage interdisciplinary research. Much of the interesting work that has come from the Organization for Tropical Studies in Costa Rica is the result of con- centration on small, well-inventoried areas of forest. Inventory must not be isolated from the other subjects under discussion in this symposium. It is significant that other speakers are covering animal-plant interactions, tropical ecosystems, and integrative approaches to the study of plant structure. Future collectors need to be more aware of the research being carried on in these and other fields, and to be ready to contribute data. The lack of pollinator data in the tropics is enormous, and both the general collector and the specialist collector can contribute much to pollination ecology by making a few observations on flower visitors, scent, etc. 1977] PRANCE—FLORISTIC INVENTORY 679 Inventory in the tropics does not just include the collection of herbarium specimens which I have emphasized in this paper. It includes inventory of pol- lination mechanisms, other insect-plant relationships, phenology, mycorrhiza, types of photosynthesis, nitrogen fixing bacteria, chromosome numbers and mor- phology (Raven, 1975), self-incompatibility mechanisms ( Bawa, 1974; Bawa & Opler, 1975), hybridization—of which we know virtually nothing in the tropics (Raven, 1976c), and many other aspects which are summed up by Farnworth & Golley (1974). Let us remember the words of Merxmiiller (1970) in reference to biosystematic work in the tropics, “A conservative today who would work on insufficient materials only, would soon be a laughingstock,” and try to improve the situation rapidly. One of the most striking facts about the tropics is that the vast majority of specimens are deposited in herbaria in temperate regions. The history of settle- ment and development has dictated the distribution of specimens, and this is now a major problem for the development of systematics and conservation in the tropics. There are very few major herbaria anywhere in the tropics, and they can easily be enumerated on two hands. They include Bogor and Singapore in Malesia; Calcutta in India; The East African Herbarium in Nairobi; The Forest Herbarium in Ibadan, Nigeria; and the Jardim Botánico and Museu Na- cional in Rio de Janeiro. There are of course fortunately a large number of smaller tropical herbaria that play an important local role; for example, there are at least 49 herbaria in Brazil alone, 16 in Colombia, etc. (see Table 9). Their work is hampered by the lack of type specimens and literature. Not only are the specimens deposited in Europe and North America, but also the best literature about an area is often in a foreign language such as English or German. is lack of resources has also been accompanied by a lack of trained per- sonnel in tropical countries which has also hampered the progress of inventory. These facts, coupled with the increase of nationalism, have led to the implemen- tation of strict rules to govern collecting activity by foreigners with the result that there are some tropical areas where it is impossible for foreigners to collect at present. In order to complete the inventory of the tropics it is necessary to stimulate more training of local resident botanists (Prance, 1975), to deposit properly identified material in all tropical herbaria equipped to house them, and to pub- lish in local journals in the countries where we are working. This will not only have an effect on the standard of botany in those countries but also contribute significantly to conservation. The enthusiasm and concern for conservation dis- played by botany and ecology students in our training program in Manaus, Bra- zil, is an unforgettable experience. These young biologists will be a powerful force for conservation in a few years time. The issue of conservation is even more sensitive than collecting, and it cannot be accomplished by foreigners with- out much support from within the host country. In order to progress in the future inventory and conservation of the plant resources of the tropics, botanists must adhere more strictly to the excellent guidelines agreed upon by many major United States Research Institutions ( Hairston, 1970). 680 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 TABLE 9. A summary of South American herbaria and areas covered. Number of. Total No. of Area of Country Country Herbaria Herbarium Specimens (km?) 1 16 260,000 1,138,914 Venezuela 5 190,000 912,050 uyana 2 30,000 214,969 Surinam 1 16,000 163,265 French Guiana 1 13,500 91,000 Ecuador 4 7,500 283,561 Peru 6 2/75,000 1,285,216 Brazil 49 2,000,000 8,511,965 Bolivia 1 1,098,581 Paraguay 1 1,000 406,752 Uruguay 3 115,000 177,508 hile 4 0, 756,945 Argentina 22 3,000,000 2,776,889 TOTALS 115 6.068,50 „068,500 17,817,615 U.S.A. (1974) 1,127 45,811,608 9,360,882 Inventory of the tropics is not nearly complete, yet destruction of their natu- ral ecosystems continues not only unabated, but at a faster rate than inventory. ere is an urgent need to accelerate the process of inventory and at the same time to encourage alternatives that will buy time for us by delaying the destruc- tion of the world’s richest biome. The more knowledge we gather about the ecosystem the better the possibility that we can use it on a sustained-yield basis. In the meantime we should do all we can to encourage some of the alternatives: the exploitation of seasonal forests (Budowski, 1976; Goodland & Irwin, 1977), floodplain forests (Goodland & Irwin, 1975; Prance, in press), of secondary forest (Budowski, 1975; Farnworth & Golley, 1974), and better distribution of food produced in temperate regions. Clearly there is still an enormous challenge ahead of us in the task of a complete tropical inventory. LITERATURE CITED ApAMs, C. D. 1972. 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Styles (editors), Tropical Trees, Variation, Breeding and Conservation. Academic Press, London. 243 pp. Woopsos, R. E. & R. W. Scuery. 1943-. Flora of 8 Ann. Missouri Bot. Gard. 30: 7-403 et seq. PLANT MORPHOLOGY AND ANATOMY IN THE TROPICS— THE NEED FOR INTEGRATED APPROACHES P. B. TOMLINSON! Plant morphology, like justice, must not only be done, it must be seen to be done. It must be done because it is foundational to many major disciplines— systematics, ecology, and plant physiology. Similarly plant morphology is ini- tially an observational discipline; the pun is intentional because I want to speak from the point of view of the research worker who needs direct access to his material and the opportunity to “see” the functional significance of form and structure which is investigated. Modern high speed travel has made easy the traditional process of what may be called “body-snatching,” i.e., the initial collection of fluid-preserved or dried materials which the plant morphologist uses for much of his work. Body-snatch- ing has contributed the largest part to our understanding of tropical plants and needs to be actively encouraged because it provides the initial comparative back- ground to any biological enquiry. Here I want to emphasize the need for elab- oration of or departure from this classical approach in what I will call inte- grated studies of biological features of tropical plants. Integration here has a dual meaning, it refers to the need to combine elements of disparate disciplines— anatomy, physiology, biochemistry, ecology, and plant-animal interactions—but also emphasizes an approach which recognizes the organism itself as an inte- grated entity so that something of its total biology is revealed. Corypha provides a simple but dramatic example of a tropical plant consist- ing of a single hapaxanthic module in which the switch from vegetative to sex- ual growth is complete, with marked contrast between the massive unbranched vegetative axis to the highly branched determinate inflorescence, with resulting proliferation of another generation of meristems. Here vegetative and repro- ductive phases are sharply segregated, but they must occur in the right sequence and at the right time. A striking example, which illustrates the way in which a branched organism can function as an integrated physiological unit is provided by Cerberiopsis candelabrum (Apocynaceae), a small tree in New Caledonia which is monocarpic (Veillon, 1971). Here a tree that is architecturally pre- cisely branched in the initial vegetative stage eventually shows synchronous flowering and fruiting which ends in its death—rather like many annual weeds. A second species, C. comptonii, a treelet with smaller leaves, is not monocarpic. This genus therefore provides material for the study of a biological problem which may be approached comparatively but requires access to field popula- tions. The plants under consideration are large and can only be represented in herbaria by fragments. An appreciation of the adaptive significance of this life style would draw on several disciplines—anatomy, physiology, and reproductive biology. It has to be appreciated that modern travel in the tropics is easy and a 1 Harvard University, Harvard Forest, Petersham, Massachusetts 01366. ANN. Missourt Bor. Garp. 64: 685-693. 1977. 686 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor.. 64 relatively inexpensive research item, compared with the many requirements of modern scientific investigation such as equipment, chemical analysis, technical support, and storage facilities. Also, common organisms which are easily acces- sible need prime consideration, there is no initial need for expeditions to remote areas. We are not looking here necessarily for new approaches; great scope is provided simply by the enormous diversity of tropical plants. The greatest em- phasis needs to be given to the study of the morphology and anatomy of woody plants since trees dominate many tropical ecosystems with great floristic rich- ness. This diversity is readily documented and quantified (e.g., Poore, 1968; Ashton, 1969; Rollet, 1974; Hallé et al., 1978). Here I attempt to demonstrate by means of a few selected examples the ways in which the scope of classical plant morphology and anatomy can be broadened by emphasizing functional relations, integrating the approaches of specialists in several fields, and adopting a holistic view of the plant. The object is to illustrate examples which in turn can provide guidelines. PRIMARY MERISTEMS From the point of view of the population biologist the “individual” in a trop- ical forest may or may not be easily recognizable for demographic purposes since clonal propagation of plants makes possible a distinction between “ramet” and “genet,” to use the terms of Harper & White (1974). A useful starting point for the morphologist is to consider primary meristems as the unit making up the forest, following the suggestion of Oldeman (1974). It then becomes possible to study the ways in which primary meristems originate, function, are protected, interact with each other, and eventually die, adding a time scale and dynamic considerations to classical plant morphology. Of course, this approach is only one of many which could be adopted, but it seems of fundamental biological significance. In this approach one can make the useful distinction between seed-originat- ing meristems (i.e., via sexual reproduction) and vegetative meristems (both of which may be either latent or active). The production of seed-originated meri- stems centers on floral biology, itself an integrated discipline which will be mentioned later. Activated seed meristems are found in germinating seeds and the subject is resplendent with morphological detail (e.g., Burger, 1972; Duke, ; Ng, ] ) which now requires an extension into functional analysis. Ecology, physiology (e.g. of dormancy), population biology, and morphology are all interdependent at this critical phase, and yet there are few studies aa attempt a functional explanation of seedling characters. Jackson (1974), some extent, succeeded in his recognition of cryptogeal germination because he added field observations to his morphological data. The classical distinction between hypogeal and epigeal germination (or cryptocotylar and phanerocot- ylar, dependent on semantic tastes; Duke (1969)—which Ng (1978) has shown to be too simplistic to accommodate tropical diversity—presents in itself two contrasted biological life styles still awaiting exploration, and shows interesting analogies with contrasted branching processes (prolepsis and syllepsis), as pointed out by Hallé et al. (1978). 1977] TOMLINSON—MORPHOLOGY AND ANATOMY IN THE TROPICS 687 BRANCHING The vegetative meristems are of prime concern to the plant morphologist since they originate the structures in which he is interested. For active primary meristems, interaction between them within the individual tree have been out- lined by Hallé & Oldeman (1970) who have provided a conceptual framework which now makes it possible to talk about tree form in a comprehensible way. This foundation can be built upon by the morphologist. An important develop- mental approach is to study ways in which vegetative meristems multiply, as in the study of branching patterns. The element of discovery which is still pos- sible in descriptive tropical plant morphology is illustrated by the recent demon- stration of equal dichotomy of vegetative apical meristems in a number of angio- sperms (e.g., Boke, 1976; Fisher, 1976, Tomlinson, 1971; Tomlinson & Posluszny, 1977). That this is not a factor of any direct evolutionary significance is easily argued, especially with recent methods of mathematical analysis of branching patterns to provide a much needed background of quantification and theory (especially Oohata & Shidei, 1971). Equal dichotomy permits only a minimal value for a bifurcation ratio, whereas high bifurcation ratios seem adaptive in many ecological circumstances (Whitney, 1976). It seems reasonable to assume that the early development of highly controlled axillary branching in vascular plants has made possible the elaborated vegetative body of modern angiosper- mous trees (and, of course, other life forms), a statement which is little more than axiomatic (Tomlinson, 1978). A mathematical analysis of branch form pro- vides a useful conceptual framework. It is still interesting that gymnosperms ap- pear to lack any mechanism for generating shoots with distichous phyllotaxis and so have lost one degree of freedom much exploited by the angiosperms in their branch architecture, especially the plagiotropic shoots of many monocotyledons. Axis differentiation in woody plants, which is a major parameter of Hallé and Oldeman’s system, in the most specialized situation has a simple morpho- logical basis since shoots with contrasted orientation can have different phyllo- taxis—spiral in orthotropic, distichous in plagiotropic. The ability of the same genotype to support such contrasted primary meristems which operate contem- poraneously provides scope for extended morphogenetic analysis. A comparable example is found in shoots with phase change from one type of orientation to the other during the activity of a single meristem. The best examples are provided by rhizomatous herbs where foliar dimorphism associated with sympodial growth is common, as in many Zingiberales. Syllepsis, as contrasted with prolepsis (using these terms in the expanded definition given by Tomlinson & Gill, 1973) provides the most convincing dem- onstration of how limiting of elementary concepts temperate-based botany may be, since syllepsis (development of a lateral axis contemporaneous with its par- ent) is common only in tropical woody plants and is little developed in north temperate woody plants because lateral axes here usually undergo a period of dormancy before extension (i.e., show prolepsis). And yet these terms are needed to describe two fundamentally contrasted conditions. Why syllepsis should be so characteristic of tropical trees still awaits an ecological explanation, but recog- 688 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 nition of this simple developmental distinction opens up numerous opportunities for comparative anatomy. The combination of anatomy and translocation physiology which integrates this dynamic aspect of shoot morphology is likely to be found in the recent dem- onstration by Zimmermann (1978a, 1978b) of the unequal distribution of hy- draulic conductivities in trees. In particular, there are pronounced constrictions at every branch-trunk junction. This must have an anatomical basis and perhaps even provide a causal explanation for continued apical control of one type of shoot over another. Since prolepsis and syllepsis may determine differences in axis orientation in many trees, there is the possibility of analyzing continuity between dynamic morphology and subsequent function in a novel way. anatomy of branch insertion may thus become as important in understanding the ecological significance of form in woody plants, as has comparative study of the stem-node-leaf continuum provided material for systematic and evolutionary analysis ( Howard, 4). Secondary changes in axis orientation depending on differences in vigor that suggest hormonal mechanisms of control are important in a number of tropical trees (Koriba’s model). Reaction anatomy as a functional mechanism in the organization of woody plants remains little explored, but is a topic likely to be a rich source of information in the future (cf. Fisher, 1978; Tomlinson, 1978). BUD BIOLOGY The persistence or otherwise of vegetative meristems in plants depends not only on genetic organization but also, as determined by ecological accident, on the efficiency with which shoots and their associated primordia are protected, as from predators, drought, or excess heat. Devices which can be interpreted as protective are often mechanical and conspicuous, but biochemical devices are probably equally if not more common. Here developmental anatomy, morphol- ogy, and organic chemistry need to be integrated. For larger shoots, size alone, combined with the rigidity of mature appendages, may be sufficient, as in woody monocotyledons. Many palms have efficient supplementary organs, like spines and mechanical leaf bases, as befits plants which are vulnerable because they may possess a single apical meristem incapable of vegetative branching (Uhl & Moore, 1973; Tomlinson, 1962). Pandanus is typically protected by serrated leaf margins. The morphological diversity of buds in dicotyledonous woody plants is well known since they so often offer useful diagnostic field characters, but the only extensive summary refers largely to temperate species (Lubbock, 1899). Although the mechanical efficiency of enveloping stipules, leaf bases, and petioles is very evident, this alone does not explain why buds seem often to be the last organs to be attacked by insect predators—or why buds can survive in the absence of mechanical sheaths. The *biology of buds" is a little explored field where the morphologist will need the assistance of biochemists in order to make progress, I believe. The concept of plant apparency ( Feeny, 1976) has to be put in a morphological context, with the relative vulnerability of different parts contrasted. In making very general surveys of tropical plants one can, for example, dis- 1977] TOMLINSON—MORPHOLOGY AND ANATOMY IN THE TROPICS 689 tinguish “wet” buds from “dry” buds, the former characterized by some fluid or resinous secretion. Wet buds commonly are associated with stipular devices which support colleters or equivalent glands, as in Rubiaceae, Rhizophoraceae, and Polygonaceae. Here the stipule may simply provide the cavity which ac- commodates the fluid secretion. Even where the stipules are small ( “vestigial” to some comparative anatomists), as in many Euphorbiaceae, Ulmaceae, Celas- traceae, and Elaeocarpaceae, they are likely to perform a vital function, since they can mature before associated leaf primordia, and their biochemical spe- cialization may be indicated by their high tannin content, an observable micro- scopic feature. In other buds which lack stipules there is no such division of labor, and the leaf primordia themselves are tanniniferous. Secretions which dry as conspicuous, resinous or varnishlike coatings of unknown chemical composi- tion, but probably polysaccharides, are common in tropical woody plants—they may make the bud distasteful, resistant to drying, reflect damaging wave-lengths, and mechanically impede chewing insects. This exuded material is something that production ecologists should not overlook, since it is often exfoliated in considerable quantities. In Ceriops ( Rhizophoraceae) between 25 and 40% of the dry weight of the bud is made up of this varnish. Initially it simply fills the quite considerable free space between stipules and leaf primordia—at this stage it is a close packing device which is related to the periodicity of growth extension; subsequently the varnish becomes a casing to the expanding leaves and inter- nodes, with unknown biological properties; finally it sloughs off. The material has not yet been analyzed biochemically, but a large amount is produced and lost each time a leaf pair expands. A possible biological function for bud secretions in other Rhizophoraceae has been shown recently by Richard Primack, in Queensland. He found that a galactose-rich exudation served as a bird-attracting device in Rhizophora stylosa, since nectar-feeding birds would lick this sweet fluid which is apparently pro- duced by the nonvasculated stipular glands (colleters). Such birds are likely to also pick off insects, this grooming being of presumed benefit to Rhizophora. Here, therefore, initial studies on the anatomy of colleters have an extension into biochemistry and plant- animal interactions. e term "naked bud," which is sometimes used to describe meristems with- out specialized mechanical protection, is not particularly appropriate because leaf primordia at an arrested state of development are frequently associated with a specialized but ephemeral indumentum, or with latexlike secretions. The ini- tial need is not for elaborate categorization, but for the examination of specific case histories with the concept of the bud as a biological unit given prime con- sideration. Emphasis must be given to the construction of buds in relation to the method of shoot extension, i.e., whether rhythmic, continuous, or intermittent (without regular periodicity). In many tropical woody plants a stipular organ can serve as an "instant" bud scale, the shoot ceasing to extend without reference to any endogenous rhythm but still retaining a protective cap. The relation between stipule position and the region of extension is of interest. In most examples internodal extension occurs beyond the level of stipule insertion, but in some 690 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Cunoniaceae, a family characterized by well-developed interpetiolar stipules, the stipule pair is carried up with the bud by extension of the internode below it. The idea that bud morphology is a dynamic and not a passive subject is one that can be encouraged by the comparative study of the wide diversity of plants available in the tropics. Internal secretions, most noticeable in those numerous tropical families with latex, often of commercial importance (Apocynaceae, Asclepiadaceae, Moraceae, Sapotaceae, and Euphorbiaceae) provide an area for integrated studies com- bining anatomy, developmental morphology, chemistry, and adaptive biology. This can be illustrated in a spectacular way by the recent report that the New Caledonian endemic Sebertia acuminata (Sapotaceae) accumulates as much as 25% by dry weight of the heavy metal nickel in its latex (Jaffré et al., 1976). This is undoubtedly an exceptional case, but it does indicate part of the mecha- nism whereby a plant can tolerate soils with a high content of heavy metals and shows an interesting correlation between plant anatomy and mineral nutrition. Both the distribution of laticifers within this plant, and the distribution of heavy metals and inorganic compounds in laticiferous organisms offer themselves as subjects for study. It is known that nickel accumulators are not necessarily lati- ciferous (Brooks et al., 1974), but where does nickel accumulate in plants of high nickel content? Electron microprobe analysis of appropriate organisms could usefully integrate ecology, anatomy, and chemistry. Of interest is the way in which herbarium specimens have been used in this research (Brooks et al., 1977) showing the lasting value of “snatched bodies.” FLORAL BIOLOGY This area represents perhaps the most profitable one for integrative studies, and a number of recent workers have combined comparative and developmental anatomy with field study of flower visitors and pollination biology. In part this is a response to Carlquist’s (1969) critique of the general subject of floral anat- omy, but much reflects the increasing field orientation of modern morphologists. The most extended study of this kind is that of Uhl & Moore (1977) on palms in which a syndrome of characters is described in detail for inflorescence and flower patterns in six examples representing two anemophilous and four differ- ent entomophilous modes of pollination. This study has as its basis one of the most complete systematic and anatomical backgrounds known for any family of tropical plants (Moore, 1973) and should serve as a model for future studies. It extends the concept of “protection” of meristems to ovules and pollen, but back- ward in time to the vegetative meristems which ultimately produce them, via the often elaborate inflorescence to the floral envelopes and mechanisms which assist in fertilization and then anticipates the later processes which contribute to the formation of seeds and fruits. We therefore add a time scale and additional biological dimensions to comparative studies which began with studies of floral vasculature. Reproduction “strategy” may even have to be integrated with photosynthetic “strategy” since position of flowers or inflorescences can determine the architec- tural model in Hallé and Oldeman’s system. Of interest are those contrasted 1977] TOMLINSON—MORPHOLOGY AND ANATOMY IN THE TROPICS 691 examples where the presence of terminal versus lateral inflorescences, resulting in determinate or indeterminate axes, in turn produces an architecture which is the morphological analogue of monolayer and multilayer, to use the terminology of Horn (1971) for probable contrasted photosynthetic strategies in trees. This kind of example shows how emphasis of a particular dynamic aspect of one phase in the life cycle of a plant inevitably leads from one topic to another, such is the nature of an organism as an integrated whole. A field-orientated approach to comparative morphology which illuminates floral structure is shown by recent studies on the mangrove Rhizophoraceae (Tomlinson et al., in preparation). This involves 4 genera and about 20 species with a comparable vegetative morphology (architecture, bud morphology) and habit (marine swamps) but with evident niche diversification shown by quite complex ecological zonation, which is reflected morphologically in diversity of aerial root development. A common floral plan is involved, indicative of a com- mon evolutionary ancestry but with immediately obvious variation in such fea- tures as inflorescence branching, flower size, orientation, and number of parts. A dominant feature is the dehiscence of stamens within the unopened flower. Functionally at least 6 types of floral mechanism can be recognized according to the way parts behave in relation to pollen vectors. These types transcend taxonomic boundaries because a single genus can include contrasted mechanisms (Ceriops) or be adapted to different visitors (Bruguiera). An unusual mecha- nism which is the result of considerable developmental complexity is a catapult release of pollen, with stamens initially enclosed by petals in a spring device triggered by a flower visitor. Despite this common piece of engineering, flowers are visited more or less exclusively by birds on the one hand, insects on the other, with further specialization according to the type of insect—moth versus butterfly, for example. Extending these observations in a comparative way, considerable circum- stantial evidence accrues that the genus Rhizophora is an exceptional component of mangrove communities because it is wind pollinated. This may be one reason for its evident ecological success and is the basis for an understanding of genetic aspects of its species interrelationships which are suggested by certain taxo- nomic peculiarities in the genus. This step-by-step development of our under- standing of this widespread and important genus began with simple curiosity about its morphology and anatomy. The essential ingredients in the success of this continuing research have been repeated access to natural populations and collaborative work with other specialists—field ecologists, population biologists, and biochemists and attempts to understand different components in a biological continuum. CONCLUSIONS The examples chosen are few and refer to a restricted field. Similar ap- proaches to different topics could have been adopted. Instead of primary meri- stems, the secondary meristems of tropical woody plants could have provided a focal point. Knowledge of fluctuations in the activity of vascular cambia in tropical trees is very scant and we largely lack the important ecological param- 692 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 eter this provides in temperate trees—the ability of an observer to determine tree age quite accurately is missing in tropical forests. The structure and development of root systems in tropical plants is little explored, especially the interaction between roots and soil microorganisms (cf. Janos, 1975). For this topic, even descriptive morphology is at a very elemen- tary stage. One particularly useful field which needs expanding is the study of the morphology and anatomy of tropical crop plants since a knowledge of their response to pathogens and pests depends on a knowledge of their normal struc- ture. However, there are no detailed and comprehensive accounts of the struc- ture of major tropical crop plants like coconut, oil-palm, coffee, cocoa, rubber, and so on. A particular deficiency is in studies of development morphology. The integrated activities of a diversity of workers is required here. The conclusion then is that plant anatomy and morphology remains a cen- tral field of tropical inquiry, but not as an isolated or static discipline. The mor- phologist has to combine his specialized abilities with those of colleagues in other fields. Once this elementary principle is accepted we can move on to the more important task—devising the most efficient means to apply this principle. LITERATURE CITED AsHTON, P. S. 1969. Speciation among tropical forest trees: some deductions in the light 96. of recent evidence. Biol. inn. Soc. 1: 155-1 Boxe, N. H. 1976. Dichotomous branching in Mammillaria (Cactaceae). Amer. J. Bot. 3: —1384. Brooks, R. R., J. LEE & T. JAFFRÉ. 1974. Some New Zealand and New Caledonian plant accumulators nickel. J. Ecol. 62: 493-499. Reeves & T. JarrnÉ. 1977. Detection of nickeliferous rocks by analysis of ia specimens of indicator plants. J. Geochem. Explor. 7: 49-57. EE: D. (HZN). 1972. Seedlings of Some Tropical Trees and Shrubs mainly of S. E. entre for Agricultural 8 and Documentation, Wageningen, Netherlands. C Ce S. 1969. Toward acceptable evolutionary interpretations of floral anatomy. s. 19: 332-362. Dure, J. A. 1969. On tropical tree oe I. Seeds, seedlings, systems and systematics. n 1 Bot. Gard. 56: 125-1 FEENv, P. 1976. Plant apparency and E defense. Rec. Adv. Phytochem. 10: Fisugn, J. B. 1976. marr of dichotomous branching and axillary buds in omen ( Monocotyledoneae ). d. J. Bot. 54: 578- . 1978. A vati erm study of Terminalia- bmnching: Chap. 13, in P. B. Tomlin- son & M. H. Sr lea (editors), Tropical Trees as Living Systems. Cu is Univ. ork. HALLE, F. & R. X A. OLDEMAN. 1970. Essai sur Architecture et la Dynamique de Crois- sance des Arbres Tropicaux. Masson et Cie, Paris. — A. OLDEMAN & P. B. ToMLINsoN. 1978. Tropical Trees and Forests—an Architectural Analysis. Springer Verlag, New York. Harper, J. L. & J. Wurre. 1974. The demography of plants. Annual Rev. Ecol. Syst. 5: 419 Horn, H. S. 1971. The Adaptive Geometry of Trees. Princeton Univ. Press, Princeton, Je Y Howanp, R. 1974. The stem-node-leaf continuum of the Dicotyledoneae. J. Arnold Arbor. 585 125-181. Jackson, C. 1974. Cryptogeal germination and other seedling adaptations to the E of vegetation in savanna regions in the origin of the pyrophytic habit. New Phytol. 780. JAFFRÉ, T., R. Brooks, J. LEE & R. D. Reeves. 1976. Sebertia acuminata: a hyperac- cumulator a nickel from New gode Science 193: 579-580. 1977] TOMLINSON—MORPHOLOGY AND ANATOMY IN THE TROPICS 693 Janos, D. P. 1975. Effects of vesicular arbuscular mycorrhizae on lowland tropical rain forest trees. Pp. 437-446, in F. E. Sanders, B. Mosse & P. B. Tinker (editors), Endom orrhizas. Academic Press, London Lussock, J. 1899. On Buds and Stipules. Kegan Paul, Trench, Trübner & Co., London. Moo . E. 1973. The major groups of palms and their distribution. Gentes Herb. 11: Nc, F. S. P. 1975. The fruits, seeds and seedlings of Malayan trees. I-XI. Malaysian For- 3-98. 1978. Strategies of establishment in Malayan forest trees. Chap. 5, in P. B. Tom- mb. & M. H. Zimmermann (editors), Tropical Trees as Living Systems. Cambridge Univ. Press, New York. OrpEMAN, R. A. A. 1974. L'architecture de la forêt guyanaise. Mem. O.R.S.T.O.M. 73. Oonata, S. & T. Sumer. 1971. Studies = 155 1 yas structure of trees. I. Bifurcation ratio of trees in Horton's law. Jap. J. E Poore, M. E. D. 1968. Studies in 1 rain pm I. The forest on Triassic sedi- ments in Jengka Forest Reserve. J. Ecol. 56: 143-169. RoLLET, B. 1974. L'Architecture ve Foréts Denses Humides Sempervirentes de Plaine. C. T. F. T. je s -Marne, Franc TOMLINSON, P. B. 1962. The leaf A in palms. Its morphology and mechanical biology. an Arbor. 43 —46. The “hagi apex and its dichotomous branching in the Nypa palm. Ann. Bot. (London) 35: 865-879. Branching and axis differentiation in tropical trees. Chap. 7, in P. B. Tom- nA: & M. H. Zimmermann (editors), Tropical Trees as Living Systems. Cambridge Univ. Ws New York — . GL. 1973. Growth habits of tropical trees: some guiding principles. Pp 129- D n B. J. Meggers, E. S. Ayensu & W. D. Duckworth (editors), Tropical Forest 5 in Africa and South America: A Comparative Review. Smithsonian Institu- tion Press, 5 D. U. USZ NY. 1977. Apical dichotomy demonstrated in the angiosperm Flagel- laria. Science 196: 1111-1112. B. Primack & J. S. Bunt. Submitte a to fo Floral morphology in rela- tion to floral biology in mangrove Rhizophor Unt, N. W. & H. E. Moore. 1973. The a of pollen and ovules in palms. Principes 17: 111-149. & 977. Correlations of inflorescence, flower structure, and floral anatomy with pollination in some palms. Biotropica 9: 170-19 VEILLON, J.-N 1. Une Apocynacée monocarpique a Nouvelle-Calédonie Cerberiopsis 5 Vieill. Adansonia, sér. 2, 11: 625-639. Wuirney, G. G. 1976. The bifurcation ratio as an indicator of adaptive strategy in w oody plant species. Bull. Torrey Bot. Club 103: 67-72. ZIMMERMANN, M. H. 1978a. Structural requirements for optimal water conduction in tree stems. Chap. 22, in P. B. Tomlinson & M. H. Zimmermann (editors), Tropical Trees as Living Systems. Cambridge Univ. Press, New Yor — —. 1978b. Hydraulic architecture dE some diffuse porous trees. In preparation. A CONTRIBUTION OF RAIN FOREST RESEARCH TO EVOLUTIONARY THEORY"? P. S. ASHTON? Though by no means universal even in the lowlands, species diversity within a single life-form reaches unequalled levels in many tropical forests, and in par- ticular in the aseasonal wet oceanic climates of the Far East. There large genera, many of whose species may occur together and are apparently spatially inter- changeable (e.g. Poore, 1968) are particularly frequent and have prompted speculation as to their origin. Ecologists (Poore) and taxonomists (Fedorov, 1966; van Steenis, 1969) alike have concluded that chance events are the major determinants of survival and must hence influence the course of evolution. Jan- zen's (1970) attractive theory that interactions between host-specific predators and their tree prey provide a density controlling mechanism which allows accre- tion of floristic diversity has yet to be investigated within large tree genera and does not apply within the Dipterocarpaceae, dominant trees of the Far Eastern rain forest canopy, whose predators are well known and are not specific even at generic level. How old are tropical tree species? How niche specific are they? Is evolution continuing within these forests? Are these communities in evolutionary i ed rium, following a long period of gradual stabilization (e.g., Stebbins 1974), « does species diversity continue to increase? What are the component tree spe- cies, are they outbreeders and are they genetically variable, or are they genet- ically uniform, even apomictic? Richards (1963) has reasoned that ecology cannot afford to ignore the tropics; an understanding of the evolutionary biology of this most species-rich vegetation must be accepted as equally essential if only to put, by comparison, knowledge of our younger and less diversified temperate counterparts into truer perspec- tive. Studies in the Dipterocarpaceae and their forests over the last 20 years, in which I have collaborated, are beginning to elucidate this subject. Tur AGE or WEST MALESIAN Forest ECOSYSTEMS Haile et al. (1977) have established that the Malay Peninsula and southwest Borneo have remained within 20° N. latitude of the equator since the late Cre- have described the work of successful and happy ee in which I have been one at many participants: Paul Chai, Ilias Pa'ie, Othman Haron and Caroline Taylor in par- ticular played a major part in the ecological research in Borneo. En gkik Soepadmo codirects the research on breeding systems which is being carried out by S. Appanah, Chan H. T., Gan V., Ha C. O., A. Kaur, and Yap S. K. under the supervision of J. I. Furtado, K. Jong, D. W. Lee, A. C. Marshall, J. D. Matthews, N. Prakash, F. W. Robertson and V. E. Sands at the Universities of Malaya and ipsas this is supported by grants from the Leverhulme Trust Fund, the Royal Society of London, the U. K. Natural Environment Research Council and the Carnegie Trust for the Universities 3 Se xland. * The organizers of the Symposium are grateful to the Atkins Garden Fund of Harvard University for funds which made possible the pa rticipation of P. S. Ashton. “Institute of S. E. Asian Biology, University of Aberdeen, Scotland. ANN. Missouni Bor. Garp. 64: 694-705. 1977. 1977] ASHTON—RAIN FOREST RESEARCH 695 taceous. Muller (1968, 1972) has described the transition from gymnosperm for- est to predominantly angiosperm forest during and following the Cenomanian, and the successive accretion of new orders and families through the Tertiary, from pollen analysis of the Plateau Sandstone formation of northwest Borneo. By the end of the Tertiary the inland forest flora was apparently not dissimilar to that of the present day, fossil leaves of putative Pliocene age from Manila for instance (Merrill, 1923) being identifiable with species still growing nearby. The pollen record suggests, on the one hand, that the region has remained within the humid, though not necessarily aseasonal, tropics since the origin of angiosperm forest, and, on the other, that the growth of floristic diversity at ordinal and familial level has occurred through a sequence of periodic and rather sudden immigrations, rather than through gradual evolution in situ. The Quaternary era must inevitably have witnessed the invasion of seasonal rainfall regimes during the periods of custatic fall in sea level, when the Sunda Shelf region, comprising the Malay Peninsula, Sumatra, Java, Borneo and the intervening seas, became a continent comparable in size and latitude to the northern part of South America; the last such time ended ca. 15,000 years B.P. The only evidence for climatic change in the lowlands during the Pleistocene is indirect, through the existence and gradual extinction of the nonforest large mammal and essentially Asiatic Trinil fauna during mid-Pleistocene times in Java (Medway, 1972). Evidence for the existence of periods of rainfall season- ality from other parts of lowland Sundaland during the Pleistocene is presently lacking. It is nevertheless difficult to believe that the extraordinary species rich- ness of west Malesian forests, which is strongly restricted to the aseasonal region south of the Kangar-Pattani line in the Peninsula (Whitmore, 1975 )—though generic diversity is not restricted to the same extent—has arisen in the last 15,000 years; the exceptionally high level of local endemism in some coastal regions, notably east and northwest Malaya and along the northwest coast of Borneo, almost certainly has a more ancient origin. Indeed, whereas localized centers of diversity and endemism exist in a sea of uniformity in South American forest ecosystems, and are considered to indicate the sites of refugia for rain forest species during the interpluvials, the greater part of the present west Ma- lesian archipelago must be regarded as analogous to one vast refugium frag- mented by the current high sea level. RATE or SPECIATION WITHIN WEST MALESIAN TREES Muller (1964, 1972) has identified the pollen type of the monotypic riparian palm Nypa from as early as the Upper Cretaceous, and in another unique paly- nological study traced evolution within the mangrove genus Sonneratia and its fossil progenitor Florschuetzia since the Eocene. Plant distributions provide indirect evidence for continuing speciation during the Pleistocene and up to the present day; in this respect dipterocarps are particularly apt subjects owing to the absence of any known fruit vector. This may be exemplified by comparing distributional patterns in Dipterocarpaceae with the postulated Pleistocene geo- morphological history of the northwest Borneo neogeosyncline (Ashton, 1972; Wall, 1967); the history of river capture neverthless remains to be confirmed by 696 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 108 are widespread, but:- 57 are confined E. Only 7 are confined 27 are confined 3 species are confined E, or W, of or W. of the ancient by the Rejang, by the insigni- the Suai-Sibuti drainage, which river Lupar, whose largest river in ficant Kemena, follow the putative course during northern Borneo; which neverthe- Pleistocene valley of the Pleistocene periods its present course less follows Baram, a major river of continentality is seems to have the putative which limits no marked by submarine resulted from late former Rejang species range. canyons northwards Pleistocene river valley. towards the conti- capture. e o š \ 3 apt nental shelf. I < = k N / ` SCA ' " ' F 4^ — Ficung 1. Geographical barriers and the distribution of 201 dipterocarps in northwest Borneo. The southeast to northwest trending drainage of northern Borneo is of probably late Pliocene-early Pleistocene origin. analysis of river sediments. Though the major discontinuity in dipterocarp dis- tributions follows a river valley of probably pre-Pleistocene origin, significant discontinuities also occur across valleys of lesser age, including examples of allo- patric subspeciation (Figs. 1-2). In some cases species may be remarkably recent in origin: The ten species of Shorea sect. Pachycarpae are endemic to Borneo, surprising in itself in view of its intermittent connection with the rest of Sundaland until the Holocene. Of these only one, S. mecistopteryx Ridl. is invariably morphologically clearly de- fined and at the same time widespread. The other widespread species, S. pinanga Scheff., S. amplexicaulis Ashton, S. beccariana Burck, and S. macrophylla (de Vr.) Ashton, are very variable and morphologically intermediate forms occur fre- quently in certain restricted localities. Of the remainder, at least two, S. prae- stans Ashton and S. rotundifolia Ashton, seem by their extremely local distribu- tion, as well as by their morphological relationships with others, to be of very recent origin; the latter for instance occurs sympatrically, even side by side, with S. amplexicaulis with which it appears to be closely related. CAUSES OF SPECIATION: AN ECOLOGICAL VIEWPOINT Nevertheless, S. rotundifolia and the species in its section are exceptional among dipterocarps: Most species of the humid tropics are clearly defined; ASHTON—RAIN FOREST RESEARCH 697 1977] be, ^ cepe stellatus. o n . . R a @ssp. [e] ' Dipterocarpus geniculatus: ° Ò ssp. I o ` ssp. geniculatus A t ° : o ssp. grandis ° 4 I ° r o / o J P4 o € e° Pd amo © : e K ' O M e° ) , M L4 LI ` » A E @ posi | : ~ 7 f `... y 4, -3 -. e à he — - ` ° + hts 2 LS $ ax * a EN 3 el! ZU PLI 2 ^» 1. East and West of the Lupar 2. East and West of the Kemena AA Shorea macroptera. ° 7 © 1 OSSp. mac ey h @ssp. bai nl , r ! J @botn eso pem ier 166 wla ° L— Y Ecotypic sympatric diversification 3. Migration subsequent to allopatric diversification Ficure 2. Patterns of speciation in dipterocarps in northwest Borneo closely related taxa are as a rule ecotypically differentiated in relation to site (Ashton, 1964, 1969), habit (Ashton, 1969), or physiology, as is the case with the commoner sympatric species of Shorea sect. Muticae (Symington, 1943). It is at the generic and familial level that taxa appear to be ecologically comple- mentary, and it may be at this level that the importance of predator-prey interac- tions play a major part in the maintenance of diversity. This is yet to be studied, but it would be surprising if successful speciation occurred among rain forest trees in response to predators whose life cycle is likely to be at least one hundred times shorter. Figure 3 shows that it is the extraordinary diversity at species level which distinguishes Malesian forests from all others. The sample from mixed forests in Surinam (calculated from Schulz, 1960, and probably a slight underestimate as species in some genera were not all distinguished) produced the same genus to species ratio as that from the isolated relict forests of southwest Sri Lanka, not- withstanding the higher number of genera in the former, a reflection of its conti- nental location. Pasoh forest has one of the lowest ratios calculated for Malesian Mixed Dipterocarp forest (e.g., see Ashton, 1976a). Mixed Dipterocarp forest differs considerably between sites, both in species 698 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 Species o o 1 Percentage of N N IddHd Z r3 j“ 1 6 7 8 9 0 12 Minimum Number of Species per Genus 16 18 20 22 FicurE 3. The percentage of species in genera of different sizes : samples a her mixed lowland rain forest. Solid columns: Pasoh Forest, Malaysia (2 x 5 ha; 191 genera; 484 species; ratio 1:2.5). Empty columns: Sri Lanka (3 x 2.5 ha; 98 genera; 166 s 2 1 5 ratio 1: 1.7). Hatched columns: Mapane, Surinam (5.6 ha; 152 genera; 255 species; ratio 1:1.7) . richness and in degree of floristic spatial variability; and both seem to be influ- enced by soil nutrient status. Our work in northwest Borneo indicates that spa- tial variation is measurably correlated with soil nutrients only where fertility is low; total phosphorus and exchangeable potassium become increasingly corre- lated when phosphorus levels are below ca. 200 p.p.m. (Fig. 4; Ashton & Brunig, 1975; Ashton, in preparation). Intrinsic floristic richness appears to be greatest where exchangeable potassium is between 1,000-2,500 p.p.m. (Fig. 5). Here a species/individual curve for a Heath forest site is compared with six others, selected to exemplify a general trend among 18 sites in Malaysian Mixed Dip- terocarp forest where I have carried out quantitative studies. The Pasoh curve is representative of the Mixed Dipterocarp forest of the Malay Peninsula, noted for its floristic uniformity (Wong & Whitmore, 1970, though see also Ashton, 1976c), growing in a region of Lower Palaeozoic rocks that are the oldest in west Malesia, and on an ancient land surface that has remained above sea at least since the Cretaceous. The plot was sited on Pleistocene raised riverain alluvium to ensure uniformity; forest on adjacent hillside in the event produced a very similar species/area curve. The Arip and Mersing sites each represent small islands, in neither case exceeding 50 km?, of an unusual substrate in the sedimen- tary rocks of the geologically and geomorphologically young Neogene basin of northwest Borneo; the former in fact consists merely of a narrow ridge, rarely exceeding 1.6 km wide. Bukit Lambir, near the youngest part of that basin, is Upper Miocene sandstone. These curves suggest then that intrinsic floristic di- 1977] ASHTON—RAIN FOREST RESEARCH 699 RE Principal components ordinations of 0.2 ha plots in Mixed Dipterocarp forest: Values for exchange able K (in p.p.m.) are superimposed, Left: Bukit Iju, Arip (rhyolite). Right: Bukit Mersing, Arap (basalt). versity within this region of probable Pleistocene climatic continuity is a function neither of geological or geomorphological age, nor of the area of uniform terrain and its potential influence on diversification and extinction. They do suggest that an equilibrium may be reached, in which either no further speciation is occurring, or immigration and speciation are being balanced by extinction; and that in the absence of disturbance the level of this equilibrium is determined by soil conditions. GENETIC VARIATION IN SPECIES POPULATIONS AND ITS MAINTENANCE It is as a consequence of these discoveries that a group of collaborators in the University of Malaya and the University of Aberdeen, including both staff and research students, has been investigating the genetic variability of tree popula- tions in the mature phase of primary rain forest in Malaya, and the underlying characteristics of cytology, embryology, and reproductive biology, including the behavioral ecology of pollen and fruit vectors, associated with it. This research centered at the Pasoh Forest, is presently in progress and mention will therefore only be made to results already submitted for publication. The two principal species under study are Shorea (sect. Muticae) leprosula Miq., a common, widespread and morphologically well defined and rather uni- form emergent dipterocarp, and Xerospermum intermedium Radlk., Sapinda- ceae, an understory fruit tree with similar distribution and variability. Electro- phoretic analysis of isozymal variation of populations of both species, coupled with biometric analysis of morphological variation by Y. Y. Gan (Gan et al., 1977) suggest that both have high levels of genetic polymorphism, but that vari- 700 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ° > Arip rhyolite, Sarawak. K:2100; P: 35 ° p Bukit Lambir sandstone. :1951; P:80 2504 Pasoh Forest alluvium K 38600 P:< 300 4 Bako National Park sandstone, Sarawak K:650 ; P:28 Kuala Belalong shale, Brunei. P I 5 _—® Bukit Mersing basalt, Sarawak. K: 5000 ; P :1200 n = m u a n Badas.Brunei heath forest. Š K:< 200 ; P :< 20 * m 1004 = > z "wo xo s 40 sdo 60 760 ado edo 1000 NUMBER INDIVIDUALS Species/individual curves from Mixed Dipterocarp forest, and a Heath forest, on soils of varying fertility; trees exeeding 10cm. O. (Exchangeable Potassium, total P, indicated in p.p.m. at 60cm.) Fıcure 5. Species/individual curves from Mixed Dipterocarp forests and a Heath forest. ation in gene frequency is short range. It is inferred from this that they are out- breeders with restricted pollen and fruit dispersal; this is being confirmed by studies of pollen compatibility and reproductive and vector biology (S. K. Yap, H. T. Chan & S. Appanah, in preparation). A. Kaur and C. O. Ha (Kaur et al., 1978) find both species to be diploid, with normal embryogenesis of the Poly- onum type. Chan has also confirmed high levels of self-incompatibility in two species of Shorea sect. Pachycarpae and successfully secured fruit formation from an artificial hybridization between them. 1977] ASHTON—RAIN FOREST RESEARCH 701 These species therefore conform to the pattern expected of long-lived plants in stable environments (e.g., Stebbins, 1958) and to the prevailing trends ob- served by Bawa (1974, 1975, 1977) in similar studies in Costa Rica. Nevertheless, Shorea leprosula and Xerospermum intermedium were chosen for study for practical reasons, owing to their relatively high population densities at Pasoh: 5 per ha exceeding 10 cm diameter for the former, 10 per ha for the latter, the mean for all species being 1 per ha. Species with low population den- sities comprise the vast majority, and it might be expected that maintenance of free gene exchange may be more difficult among them. Gan et al. (1977) found a very low level of genetic polymorphism by isozyme analysis in Shorea ovalis (Korth.) Bl. ssp. sericea (Dyer) Ashton, a result which may nevertheless be an artifact caused by the fact that this species is a tetraploid (Jong & Leth- bridge, 1967). Jong (1976) reported meiotic irregularities in the same species. Chan (in Gan et al, 1977; Kaur et al., 1978) found that S. ovalis appears to be fully self-compatible, and, though lacking the close intraspecific flowering synchrony that is a characteristic of most dipterocarps, it has a more uniform than average fruiting success rate. A. Kaur (Kaur et al, 1978) has now con- firmed that apomixis occurs in this species and at least one other through adven- tive polyembryony. Apomixis is also inferred through the constant occurrence of triploidy in root squashes from several seedlings originating from a single tree, and from the production of more seedlings from a fruit than there are ovules in several others. Among 16 dipterocarp species studied quantitatively by us, 10 at least sometimes produced multiple seedlings from the normally 1-seeded fruit, though this does not confirm polyembryony as the dipterocarp ovary initially bears 6 ovules. In the four species in which several individuals were under observation the proportion of seeds producing multiple seedlings varied widely between individuals. Though it is unlikely that apomixis occurs through adven- tive polyembryony in all species producing multiple seedlings, it equally cannot be assumed that it does not occur in species in which only single embryos develop. C. O. Ha (in Kaur et al, 1978) has inferred apomixis in the dioecious understory tree Garcinia parvifolia Miq., and has inferential evidence for its occurrence in other species of Garcinia. If we accept Grants (1958) view that dioecism is itself a derived condition, then apomixis must here be regarded as evidence of extreme derivation, and an example of the advanced evolutionary levels that can occur, presumably as a result of natural selection, in the rain forest environment. Though it can hardly be claimed that these few species can adequately rep- resent the West Malesian lowland tree flora as a whole, they do conclusively demonstrate that apomixis occurs within those series of closely allied species, occurring together in the same habitat, which are such a unique character of that region. An unexpected observation is that those species whose genecology might be expected to favor allopatric diversification and distinct discontinuities in varia- tion, Shorea leprosula and Xerospermum intermedium, are remarkably uniform throughout their wide range, while every one of the dipterocarp taxa in which 702 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 apomixis is inferred or confirmed are morphologically distinct and possess closely allopatric distribution patterns. Some, such as Shorea ovalis, occur in relatively high density populations and are widespread, while others such as Hopea sub- alata Sym. are extremely local; this will form the subject of a forthcoming paper. TOWARDS A SYNTHESIS To help identify priorities for future work we suggest the following hypoth- esis to explain our observations: In the uniform physical environment and pre- dictable climate of lowland Malaya natural selection will be dominated by biotic factors. These factors will continuously change in time through the periodic accretion of immigrant species into the forest community, each newcomer thus inevitably modifying the competitive interactions of those already present. The maintenance of genetic variability within species is therefore essential to their long-term survival, and maintenance of cross-pollination is at a premium. As the density of the forest community as a whole cannot increase, it must follow that the arrival of new species, and particularly those which successfully build up relatively high population densities must lead to a compensatory decline in the population density of those already present. This will hasten the decline of species whose densities are already low by increasing the physical difficulty of cross-pollination from increasingly distant pollen sources as Fedorov predicted; this in turn will lower fruit yield and weaken the reproductive pressure required for maintenance of numbers. Natural selection in species with very low popula- tion densities—and these could constitute the majority—might well favor geno- types that are both well adapted and can maintain reliable and high fruit pro- duction. It is suggested therefore that apomixis has originated in rain forest trees once again as a means of overcoming sterility, as suggested long ago by Darlington (1939 We might infer, from the variability between trees in the proportion of mul- tiple seedlings produced, that apomixis occurs in only a proportion of trees in some species populations. It is difficult nevertheless to see how a balance be- tween the number of apomictic and/or self-compatible, and obligate outcrossing individuals can be maintained in perpetuity, for increasing rarity due to chang- ing interspecific competition would lead to increasing decline in the proportion of obligate outcrossing individuals. Conversely obligate apomixis, or gene fixa- tion in small isolated populations of self-compatible individuals, must itself be regarded as an evolutionary dead end, preluding inevitable extinction in a con- tinuously changing biotic environment. Thus the maintenance of low levels of self-compatibility, and sufficiently high population densities to ensure adequate reproductive pressure to maintain numbers through outcrossing are both essen- tial adjuncts to long-term survival of a species. If obligate apomixis does fre- quently occur (and this requires much more study) and if our hypothesis is correct, we may see in the ancient Malayan rain forest a phenomenon whic must eventually arise in all plant communities: there must be an ultimate limit to the level of intrinsic species diversity that can be attained, beyond which accretion is balanced by extinction (see also discussion in Whitmore, 1975 We have here an analogue of G. G. Simpson's (1953) "evolutionary episode" 1977] ASHTON—RAIN FOREST RESEARCH 703 in the context of a multispecies community. Invading species, we suggest, are mainly outbreeders, but the low densities of even the commonest species com- bine with limited pollen and fruit dispersal to favor rapid allopatric speciation; such species will prevail in young forest communities and this can be tested in the isolated forests of volcanic islands or the Atlantic foothills of the Central American Cordillera. When numbers decline through competition, and as the overall floristic diversity increases, selection will increasingly favor apomixis which may then be an agent of secondary, essentially ephemeral and possibly sympatric, speciation. Thus the declining species do not fade away gradually but, by borrowing time in a Faustian pact of apomixis, regain the stage from time to time before their inevitable nemesis. CONCLUSIONS We now have growing evidence then that Malesian trees of the mature phase of primary rain forest are highly niche specific; that some may be old, but that speciation is actively continuing among many others; that there is a maximum number of species that a forest can accommodate, that this varies with site con- ditions and that it has already been approached in west Malesian forests; and that a remarkable variety of breeding systems exists even within the mature phase alone. An overall picture is thus beginning to emerge in which each part of the puzzle is becoming interlocked, but it hardly confirms Stebbins's (1974) picture of the tropical rain forest as merely a repository for botanical antiques! But what of the gap phase species—the woody pioneers? How have so many archaic forms continued to survive nevertheless? And what are the mechanisms that allow species diversity to reach its highest level on relatively infertile soils? A student of ours is about to embark on a study of the first, and we are planning to pursue the others in the near future. But our forest of ignorance is deep and vast, and for all its intriguing mystery attracts far too few explorers. Once again the fascination of academic theory has dominated my presenta- tion, but what are the realities on the ground? Raven (1976) has eloquently described the demise of this unique vegetation; man's destruction in pursuit of short-term gains will lead to long-term disaster for humanity, as inevitably as apomixis may for my proud dipterocarps. Everywhere it is the same—uncon- trolled and injudicious logging practices and immigration of peoples unfamiliar with local agricultural conditions, leading to destruction of the hydrological balance of catchments, physical erosion, flooding and silting of the fertile plains, the supremacy of perennial herbaceous weeds, and the final destruction of social systems and starvation. We biologists have for too long pointed our accusing fingers elsewhere, at the politicians, financiers, even the poor peasants, anyone but ourselves. I would suggest that two of the most intractable problems are essentially scientific: The lack of critical research into the most effective means of bringing about progressive change in land use based both on scientific inno- vation and traditional practices and values; and the tendency for us scientists to go for the easy options—be it the investigation of an isolated academic problem or the development of a technique to enhance short-term profitability without regard to its wider implications—and to fear the interdisciplinary collaboration 704 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 on a broad front that alone can provide the prescriptions needed. Our little venture may claim to be a hesitant start in the right direction (Jong et al., 1973; Ashton 1976a, 1976b), for in it we combine research education with a conscious choice of species which have potential in plantations, for timber and fruit. Now we must use the knowledge we have gained to experiment in the establishment and improvement of new crops for new lands—those that were considered unex- ploitable by traditional farmers and are now all that is left. We must get these crops from the much-heralded gene pool of the forest, and this we plan too. But we will need economists, agronomists, social anthropologists, and others besides before our work can reach the stage of practical applicability. Above all, we need collaboration. LITERATURE CITED i P. S. 1964. or aur studies in the Mixed Dipterocarp forests of Brunei State. xford Forest. Mem. 1-75. 196 cia. pus nd forest trees: some deductions in the light of re- cent evidence. Biol. J. Linn. Soc. 1: 155-196. 1972 The quaternary geomorphological history of western iru and lowland forest phytogeography. Pp. 35-49, in P. Ashton & M. Ashton (editors), The Quaternary Era in Malesia. Trans. of = Second Aberdeen-Hull Symp. Malesian Ecol., Univ. Hull, Dept. Geogr., Misc. Ser. 13: 1976a. An approach ^ «the study of breeding systems, population structure and taxonomy of tropical trees. Pp. 35-42, in J. Burle B. T. Styles (editors), Tropical rees, Variation, Breeding and Conservation. Academic Press, New York. 1976 Factors affecting the development and conservation of tree genetic resources in South- East Asia. Pp. 189-198, in J. Burley & B. T. = (editor), Tropical Trees, Variation, Breeding and Conservation. 255 Press, Nev 1976c. Mixed Dipterocarp forest and its va ariation with “habitat in the Malayan lowlands: A re-evaluation at Pasoh. Malaysian Forester 39: 56-72. & E. F. Brunic. "id as variation of tropical moist forest in relation to environ- mental factors and its nce to land-use planning. Mitt. Bundesforschanstl Forst-, Holz-, coa i 1 0 109: 59- Bawa, K. 8. 1974. eeding systems of tree species of a lowland tropical community. Evo- 92. = et — 75. Dioecism in tropical forest trees. Evolution 29: 167-179 e ee biology of Cupania guatamalensis Radlk. (Sapindaceae ). — 63. 1 ; Geslo 31: DARLINGTON, C. D. 1939. The Evolution of Genetic Systems. Cambridge Univ. Press, 5 FEDOROV, A. 1966. The structure of the tropical rain forest and speciation in the humid tropics. T Ecol. 54: 1-11. Gan, Y. Y., F. W. RopEnrsosN, P. S. Asu E. SoEPpApMo & D. = LEE. 1977. Genetic m in wild populations x rain iu trees. Nature 269: Grant, V 1958. The regulation of recombination in plants. Gold Spring Harbor Symp. Quant. Biol. 23: 337-36 Haire, N. S. 1973. The recognition u subduction zones in Southeast Asia. Pp. 885- 892 in D. H. Tarling & S. K. Runcorn (editors), Implications of Continental Drift to the Earth Sciences. Aca fal Press, New York. . W. McELHINNY & I. MCDOUGALL. 1977. Palaeomagnetic data and radiometric ages from the Cretaceous of West Kalimantan (Borneo), ae significance in inter- preting geological structure. J. Geol. Soc. London 133: 133- Janzen, D. H. 1970. Herbivores and the number of seed sasa, in TT forest. Amer. Naturalist 104: 501-528. Jonc, K. 1976. Cytology of the Dipterocarpaceae. Pp. 79-84, in J. Burley & B. T. Styles E Tropical Trees, Variation, Breeding and 5 Academic Press, New Vork. & A. LETHBRIDGE. 1967. Cytological studies in the Dipterocarpaceae, 1. Chromo- 705 1977] ASHTON—RAIN FOREST RESEARCH Notes Roy. Bot. Gard. Edinburgh 27: 175- some numbers of certain Malaysian genera. . C. Strong & E. SorepapMo. 1973. Ds tropical forest: an underexploited enetic Pt of edible-fruit tree ee Proc. Symp. Biol. Res. & Nat. ia 113-121. Kaur, A., C. ( „K. Jong, V. E. SANI . T. CHAN, E. SOEPADMO & 4 8. TON. 1978. Apomixis may UR d among pies of the climax rain forest. Nature 27 1: 440—441. Mepway, Lon ia: A review. Pp. 63-83, in P. Ashton & M. Ashton (editors), The 1 Ecol., Uni dead sia Aberdeen- 9e: d Malesiar 923 Distribatinn of the D ory of the m angrove vegetation MERRILL, E. MULLER, J. ins A palynological contribution to n Borneo. Pp. anwell (editor), rase Pacific "Floras: the Pollen Story. Univ. of Hawaii Press, Honolt n 1968. Palynology of the Pedawan and Plateau Sandstone formations ( Cretaceous— Eocene) in Sarawak, Malaysia. Micropalaeontology 1 7. 1972. Palynological evidence for change in 5 5 and E tion in the Mio-Pliocene of Malesia. Pp. 6-16, in P. Ashton . Ashton (editors), T Quaternary Era in Malesia. Trans. of i Second Aberdeen- Hull Symp. "Malesia n Univ. Hull, Dept. Geogr., Misc. Ser. 1-122 .D. 19 Studies in we sian rain forest, I. The forest on Triassic sedi- co 6. Poore, M. k , ments in Jengka Forest Reserve. J. Ecol. 56: 143- Raven, P. H. 1976. The destruction of the tropics. Frontiers 40(4): 22-23. š t the tropics can contribute to ecology. col. 51: 231-241. in northern denn. Verh. Kon ScHuLz, J. P. ed. Akad. Wetensch. 53(1): 1-267. Simpson, G. G. The Major Features of Evolution. Columbia Univ. Press, New York. STEBBINS, G. L. 1958. Longevity, habitat and release o genetic variability in the higher plants. Cold. Spring Harbor Symp. Quant. Biol. 23: 365-37 197 Flowering Plants: Evolution Above the Species Level. Arnold, London 8 C. G. VAN. 1969. Plant speciation in Malesia, with special reference t the 3. theory of non-adaptive saltatory evolution. Biol. J. Linn. Soc. 1: 97-13 43. Foresters manual of 5 Malayan Forest Rec. 16. 1 with SYMINGTON, C. F. Wau, J. R. D. 1967. The quaternary . history of North special reference to the Subis Karst, Niah. Sarawak Mus. J., New Ser. 15: 30- —125. Wuirmore, T. C. 1975. Tropical Rain Forests of the Far us ue ae det 5 5 . WurrMonE. 1970. On the influence of soil properties on species dis- Malatra B. 33: 42-54. Wons, Y. K. & T. tribution in a Malayan lowland dipterocarp rain forest. PROMISING DIRECTIONS OF STUDY IN TROPICAL ANIMAL-PLANT INTERACTIONS' DANIEL H. JANZEN? Plants are not just food for animals, and animals are not just decorations on the vegetation. The world is not green. It is colored lectin, tannin, cyanide, caf- feine, aflatoxin, and canavanine. And there is a lot of cellulose thrown in to make the mix even more inedible. Animals are not ambulatory bomb calorimeters. They starve, they ache, they abort, they vomit, they remember, they die, and they evolve. Peter Raven asked me to write about promising directions in tropi- cal animal-plant interaction studies, mostly because he believes there are some. Well, it’s roughly analogous to standing in the city of London after WWII and saying, well, let’s get on with studying the promising directions in London’s architectural history. paper is about tropical interactions; they are the first to be extinguished by man’s onslaught and the last to be lamented. Interactions have several traits that make them especially inconspicuous (Janzen 1974a). (a). The participants, being to some degree self-sufficient, may persist well after the interaction that produced them is gone; a Scheelea rostrata palm left standing in a Costa Rican pasture will persist long after the agouti that buried its seed and the forest that gave dry season shade to its seedlings has been removed (Janzen 1971a). (b). Humans eat participants, not interactions; being relatively incompetent until quite recently, humans have by and large not generated cultural rules for the maintenance of interactions per se, but rather for the preservation of the partici- pants. (c). Humans eat only certain participants, and often atypical ones; if the interaction is to be preserved, it is not the overall interaction in which the partici- pant happens to be imbedded that is preserved, but rather that subinteraction which will generate the largest number of participants for dinner. (d). The sys- tematics and taxonomy of interactions is hopeless; most of the types have already been mutilated or destroyed and what is perhaps even worse, it is virtually im- possible to look at an interaction and know if it is largely intact. (e). You cannot collect an interaction and keep a specimen on display in a museum. (f). An inter- action has no material potential worth, as opposed to the participants which can be noticed and retained if for no other reason than the optimistic view that some day a use may be found for one of them. However, and this is a big however, I must add in the same breath that it is as examples of how things can happen that interactions are the most valuable and therefore most deserving of preser- vation. The big problem is that human wants are generally so unrepresentative of organisms in general; the specific interactions desired by humans are not ‘This paper was provoked by Peter Raven. It was supported by NSF BMS 75-14268. Portions of the manuscript were constructively criticized by J. Karr, and D. Pearson. It is dedi- cated to our grandchildren, who will inherit a devastated tropics. Francis Ng, Peter Becker, M. L. Duncan, Marina Wong, and Daljeet Singh aided extensively in the Malaysian fieldwork. * Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania 19104. ANN. Missouni Bor. Garp. 64: 706-736. 1977. 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 707 likely to be found in nature but rather will have to be hand-tailored with the desired participants. Now after that pessimistic preamble, I am still left with the task of pointing at some promising directions in the study of tropical animal-plant interactions. There are many. I take them in no particular order, and if I ignore one of par- ticular importance to you, view it as oversight and not an evaluation. Rather than preach that we should study this or that, I will simply give brief examples to draw attention to mysterious patterns, curious new hypotheses, and perplexing observations. Gilbert (1977) has, on the other hand, presented somewhat of a challenge when he stated that “It is not clear, however, that further base-level exploration would provide many new ecological or evolutionary insights, or that additional categories of interactions would be found which fall outside those major kinds that have so far been described.” I wonder. ny person seriously interested in tropical animal-plant interactions should take a week or two to read the recent symposium and review publications in this area (Van Emden, 1973; Luckner et al., 1976; Burley & Styles, 1976; Gilbert & Raven, 1975; Wallace & Mansell, 1976; Jermy, 1976; Levin, 1976; Gilbert, 1977) and browse the numerous papers on this subject in the post-1969 issues of Ecol- ogy, American Naturalist, Science, Biotropica, Oecologia, Journal of Animal Ecology, Journal of Applied Ecology, Evolution, and the Annual Review of Ecol- ogy and Systematics (among others). PLANT PRODUCTIVITY AND THE ANIMALS IN THE HABITAT At the lowland Pasoh rain forest, Negri Sembilan, Peninsular Malaysia, I cen- sused the plants in flower that were less than 3 m tall in the understory of undis- turbed forest along 3 km of narrow trail (early September, 1976). I found one orchid, one 1.5 m tall Araliaceae, one 0.5 m tall Acanthaceae ( Lepidagathis lon- gifolia), and one 1 m tall Ixora-like Rubiaceae. In the lowlands of the national park, Taman Negara, 5.4 km of rain forest trail yielded one white-flowered ginger, two Ixora-like Rubiaceae, one Acanthaceae, one unknown family, and two 10-20 cm tall Gesneriaceae with underground stems. In primary forest understory in the new Corcovado National Park (20-160 m elevation, Osa Penin- sula, southwestern Costa Rica), a trail-side survey of 4.3 km yielded 94 plants in flower of 18 species (20 November 1976). In other words, I averaged 1.3 plants in flower per kilometer in the Malayan rain forest understory and 21.9 plants in flower per kilometer of Costa Rican rain forest understory. These woefully small samples reflect accurately my general impression of the general abundance of flowers in the understory of rain forests of Peninsular Malaysia and Sarawak, as compared with those of Costa Rican rain forest of similar elevation. I was informed locally that 1976 was one of the heaviest years in memory for flower and fruit production in Peninsular Malaysia; November is the time of most reduced flower production in Costa Rican rain forest under- story (and see Frankie et al., 1974). In short, if one were to turn loose in Pasoh or Taman Negara the rain forest understory fauna of flower-visiting humming- birds, butterflies, moths, and bees found in the Corcovado, I predict that they would be dead of starvation in a few days. Furthermore, they could not survive 708 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 by moving out into secondary regeneration; Malaysian disturbed sites have a grossly lower flower abundance than any weedy wet-season vegetation that I have seen anywhere in the African or neotropical lowlands. Over the Malaysian transects mentioned above, I encountered 63 understory individuals in fruit (22 species) for an average of 7.5 per kilometer. In the Cor- covado forest, there were 345 individuals in fruit (34 species) for an average of 78.4 per kilometer. Again, the fauna of understory birds that frequently eats small fruits in neotropical rain forests would have a very rough go of it in the Malaysian forests. It is extremely interesting that after doing this and writing the above, I dis- covered Karrs (1976) statement that "about 80% of the canopy and understory tree species on Barro Colorado Island are dispersed by animals (Foster, 1973), while only about 1076 of the trees on Fogden's (1972) [Sarawak] study area were important as sources of fruits for birds." Furthermore, at the IV International Congress of Ecology in Panama, Karr ( March, 1977) noted that "The most strik- ing difference is the total lack of undergrowth frugivores in mist-net samples taken from Malaysia as compared with 25-33% of the individuals captured in undergrowth of African and Central American forest.’ I would like to propose a rather sweeping hypothesis to account for this pau- city of flowers and fruits on rain forest understory shrubs, a paucity which should have a very depressing effect on the biomass and species richness of the under- story fauna. I need first, however, to belabor you with three facts about the lowland Malaysian rain forests in which the censuses were made. (a). They are dipterocarp forests, which means that between 50 and 80% of the tree crowns in the canopy belong to species of Dipterocarpaceae. The members of this family, in Malaysia and some other tropical Asian areas, mast fruit within (and between) habitats. Thus the bulk of the flower and fruit pro- duction by better than half of the upper canopy photosynthetic machinery is pulsed at 3 to 11 year intervals. Associated with this, the animal community is sufficiently satiated by the enormous numbers of seeds that a very large number survive to the seedling and small sapling stage (Janzen, 1974b). b). Malaysian rain forests, on the Malay Peninsula or in Sarawak, are largely perched on sandy soils ranging from very old white sand deposits (such as in Bako National Park, Sarawak) to very sandy soils derived from weathering of granitic base rock that has not been inundated by the sea for an extremely long time. There is no volcanic overlay nor crust of weathering limestone on the majority of the terrain. There are many indirect measures of the relatively low ability of these soils to generate a vegetation with a high harvestable productiv- ity for other organisms: when cleared, the second-growth vegetation is very slow to refill the site (Janzen, 1974b, 1974c, and this is probably why plantation rubber is so successful on these soil types); the forest has largely remained uncut and unexploited by agrarian peoples despite their presence in the general area for many thousands of years (note that virtually all of nearby Java on volcanic soils is under agriculture); second-growth vegetation of the sites has an amaz- ingly low insect biomass as compared to that of comparable neotropical weedy sites (Janzen, 1974b); etc. 1977] JANZEN—-ANIMAL-PLANT INTERACTIONS 709 (c). There are bees, butterflies, flower-visiting birds, small fruit-eating birds, etc. present in the Malaysian rain forests. In other words, pollinators and dis- persal agents can be drawn from these groups if the ecological and evolutionary opportunity is presented. I hypothesize that the shortage of rain forest understory flowers and fruits is largely attributable to two forces operating simultaneously and synergistically. First, I hypothesize that the large pulse of dipterocarp seedlings and saplings takes up a large part of the resources that are available to neotropical under- story shrubs; the dipterocarp offspring are apparently dying in large part through competition rather than through supporting a seed-predator guild. Simulta- neously, they are analogous to an enormous and very generalist herbivore in their impact on understory shrubs. Since dipterocarp seedlings never flower or fruit, they take a large portion of the understory resources without feeding part of it back into the flower-visitor and fruit-eater guild so conspicuous in a neotropical forest. Second, I hypothesize that as the soil conditions get progressively worse, the ability to be a reproducing individual in the light-poor understory is reduced. That is to say, irrespective of the presence of the dipterocarp seedlings, if the forest canopy is held constant and the soil fertility is depressed, the biomass (number of individuals in general) and reproductive output per ha by under- story shrubs should fall (just as it would if soil fertility were held constant and the light were decreased). In other words, the rain forests of Malaysia sit on a poorer piece of real estate than do those of lowland Costa Rica, and the flower and fruit density in the understory reflects this. The animals are probably woven into this matrix more firmly than I have indicated so far. I have hypothesized that the habitat-wide masting behavior of these Dipterocarpaceae is driven at present, and was selected for in the past, by the seed predators in general (Janzen, 1974b). Further, I have argued that the lower the overall productivity of the site, the more likely it is that the animals will select for masting behavior because the less food there is for them between mast crops, the more severely they are depressed in density by masting behavior. But the scarcer they are between mast crops, the fewer understory flower and fruit crops they can (will) visit; the fewer crops they visit, the less well off will be such plants and the better off will be the dipterocarp seedlings in competi- tion with nondipterocarps. Why doesn't the system progress to where there are nothing but seedlings and saplings of overstory trees in the understory? Prob- ably because as time passes since the last mast crop, competitive and accidental deaths clear the arena for some other species of plants, and because a number of animals that visit flowers in the understory can also go elsewhere for food; many frugivores can feed on insects and other food types when understory fruits are scarce. The focus to this point has been largely on the biomass of flowers and fruits, and associated animals. However, the species richness of plants and animals should also be negatively influenced by a reduction in harvestable productivity (Janzen, 1977c). My argument involves resource partitioning and specialization on the partitions. In short, as the productivity of harvestable resources in the habitat falls, more and more resource blocks become too small to sustain a spe- 710 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 cialist. "They are then taken by a more generalized harvester or by another trophic level. In the context of the example under discussion, the number of flower-visiting species of understory birds should decline as the soil gets poorer and as the overstory becomes progressively more synchronized at supra-annual seeding. For example, in a Costa Rican rain forest there are species and morphs (often females) of hummingbirds (e.g. Phaethornis spp.) that specialize on widely scattered understory individuals in flower, and species and morphs (often males) that specialize on large clumps of flowers on forest edges (e.g., Stiles, 1975). From what I have seen of Malaysian lowland rain forest, a hummingbird would have to forage at all such sites and then some to stay in the game. Simul- taneously the species richness of seed predators in the habitat should also decline as soils become poorer and synchrony increases, since the progressively more pulsed nature of the seed resource makes it effectively scarcer in any but the very exceptional mast year. For example, in a Costa Rican rain forest there is a large standing crop of agoutis (Dasyprocta punctata) and pacas (Cuniculus paca) that live on the rather continuous input of fruits, seeds and young seed- lings (e.g., Smythe, 1970). These animals are relatively sedentary. They do not have ecological analogues in Malaysian forests, and I suspect the reason to be that in most years the seed resource is not large enough to sustain them, though in mast years it is far greater than they could ever consume before the seeds germinate. The pulsing of productivity in a rain forest can have other interesting side effects on animals. It should select for migratory or very nomadic species, which are in turn less likely to develop local regional populations than are more seden- tary species. I have argued that the wind-dispersed nature of dipterocarp seed (and that of other trees that fruit as they do, such as the legume Koompasia) is due to their specialization to the site on which their parent grew and is not involved in escape from seed predators through dispersal (Janzen, 1977d); it may also be due to an extreme shortage of biomass of frugivorous animals owing to the fact that much of the seed production by the forest is pulsed (the frugivores would be severely satiated on seeding years, just as would be the seed preda- tors). Whatever the cause, the fact that most of the canopy-level seed produc- tion is wind-dispersed eliminates a large portion of the fruit input that is an important part of the diet of many neotropical animals. For example, I doubt very much that any Malaysian forest comes anywhere close to the figures of 1.93 g of fruit per m° calculated to fall in a Panamanian rain forest by Smythe (1970). However, in closing this paragraph, I cannot help but notice that Malaysian for- ests have an exceptionally high number of species of squirrels (e.g., 19 tree squir- rels in Borneo; Davis, 1962). It is possible that squirrels are particularly good at dealing with a highly pulsed food input, as compared with the other animals that eat seeds and fruits (some in fact, are specialists on insects or vegetative parts of plants). In short, as harvestable productivity becomes progressively less available, there is no reason to expect all animal life forms to be depressed at the same rate. In fact, the elimination of some could quite reasonably result in an increase in others. The ramifications of low productivity of harvestable resources by the plant 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 711 community in an average year can produce a multitude of higher-order interac- tions. For example, in 17 days of fieldwork and travel between field sites by boat or small car, I saw a total of three raptorial birds in Peninsular Malaysia (and none in 11 days in Bako National Park, Sarawak). The area traversed was at least 480 km of urban, rural, and forest roads, 122 km of large river through farmland and forest reserve (Tembeling River on the way to and from Taman Negara), and about 50 hours of hiking in forest reserves. At least 80% of the weather was nonrainy. I should emphasize that I was not searching for raptorial birds, but rather just watching for any kind of animal. Ina similar excursion up and down the similar-sized Sanaga River in Cameroun, I took photographs of 23 birds of prey and saw at least 50 more. In Ugandan and Kenyan forest-farm- land and national parks, it is hard to find a moment on a clear day when a raptor or large avian scavenger is not in view somewhere (and see Janzen, 1976a). In Costa Rican lowland rain forests, forest-farmland mixes, and open pasturelands, raptors and/or scavengers are seen at least once every several hours, and much more often in many circumstances. The ornithological literature is not designed so as to provide material rele- vant to comments such as those above. However, a few interesting tidbits can be extracted. For example, the black or king vulture (Torgos calvus) is common throughout the northern part of the Malay Peninsula but is almost never seen in the southern half (rainforested portion) of the peninsula; the same may be said of the other peninsular vulture (Pseudogyps bengalensis) (Robinson, 1927; Med- way & Wells, 1976). As Wells put it (personal communication), there is no vul- ture (for all practical purposes) in West Malaysia. The standard explanation for the absence of vultures is Robinson's (1927) comment that “securing their food entirely by sight, it is obvious that a heavily forested country is quite unsuited to them and it is for this reason, probably, that they do not extend to the Malay Archipelago." This seems to me to be a quite inadequate explanation. As Penin- sular Malaysia has been cleared, vultures have become rarer, not more common (Robinson, 1927). Furthermore, one has to ask (1) why similarly heavily for- ested areas in other parts of the tropics sustain vultures, (2) why the forest was not cleared for agriculture and livestock long ago as it was in other parts of the tropics, and (3) why the contemporary invasion of agricultural peoples does not bring with it adequate food for vultures? In short, I hypothesize that rain forest Peninsular Malaysian and Sarawak habitats never did generate enough carrion to keep vultures in the game, and that the contemporary peoples occupying these habitats cannot raise enough livestock to generate enough spin-off carcasses for vultures to persist as the land is cleared. Central American rain forest and associ- ated natural disturbance sites, when put into multi-use agriculture and livestock husbandry, sustain conspicuous populations of three species of vultures and two caracaras (hawks that act like vultures). I doubt that the paucity of vultures or vulturelike birds in Malaysia is due to excessive hunting; however, if there is less food for them, then even small amounts of hunting can do disproportionately more damage than if there is a large resource base. I doubt that the large varanid lizards, relatively common on riverbanks and in refuse dumps where not hunted, are competitively exclud- 712 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 — ing the vulturelike birds. I saw 28 large (0.5-1 m snout-vent) Varanus along the bank of about 20 km of the Tembeling River at and below Taman Negara on one morning. Rather, I suspect that the absence of vultures allows the presence of these relatively slow scavengers; if the food is scarce and occurs at very long intervals, then a cold-blooded professional starver would be able to maintain a much higher biomass than birds. I was told by a Kuala Lumpur “pet” dealer that with water, a large varanid can live a year without food; I doubt a vulture could do the same. The hypothesis that the natural habitats of West Malaysia generate a low density of food for large carnivorous birds is also supported by the species rich- ness of falconids and accipiters. West Malaysia has 11 resident species of accipi- ters and 1 resident falcon (Medway & Wells, 1976) and is about 132,000 km? in area; Costa Rica has at least 28 resident species of accipiters and 8 resident fal- cons and is 51,000 km? in area (Slud, 1964). The tiny Costa Rican rain forest field station at Finca La Selva (6.1 km?) has at least 9 resident accipiters and 4 resident falcons (Slud, 1960). Herons, bitterns, and egrets are conspicuously scarce in fields, roadside ditches and impoundments, rice paddies, streams, marshes, and riverbanks in West Malaysia away from the sea. I did not see a single individual in the 17 day field period. More specifically, not a single one was seen along the 122 km tra- versed of the Tembeling River, despite careful search for them. These birds are conspicuous in similar habitats in Africa and Central America. On the Sanaga River trip mentioned above, I photographed 7 species and saw at least 30 indi- viduals. Such birds are a standard part of the scenery along large Central American rivers and in the kinds of habitats mentioned at the beginning of this paragraph. Inquiry of ornithologists in West Malaysia produced two useful com- ments. First, "they are absent because they don't migrate here"; well, what is wrong with West Malaysian real estate so that migrating large piscivorous birds don't use it much as overwintering grounds? Second, *these birds are conspicu- ous in areas near the sea.” For example, Medway & Wells (1976) noted that 6 of the 9 resident species of Malayan Ardeidae are associated with mangroves. If in fact West Malaysia is a poor habitat for these birds, then the mangroves and river deltas should be the best of the sites, and appear disproportionately good compared to inland areas. Again, tiny Costa Rica has 14 species of resident Ardeidae (Slud, 1964) to compare with 9 for Peninsular Malaysia (Medway & 8, ). I hypothesize that herons, bitterns, egrets (and anhinga- and cormorant-type birds) are in short supply in the West Malaysian inlands simply because the waterways dont generate enough biomass of aquatic food for them. If the surrounding terrestrial habitats generate a reduced number of insects as well, which are an important part of the diet of many ardeids, the effect would be compounded. The biomass of vascular epiphytes in the crowns of rain forest canopy- member trees at low and intermediate elevations is conspicuously lower in dip- terocarp forests than in analogous rain forest in Costa Rica, Venezuela, Colom- bia, Cameroun (Edea Forest Preserve), and Uganda (near Fort Portal). The 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 713 quantity of bare horizontal large branches in the canopy of a forest such as that at Pasoh or Taman Negara is phenomenal. I hypothesize that the cause is that the habitat generates such weak nutrient rain (bird droppings, dead insects, ant nest debris, rainwater minerals, dust, leaf and fruit litter, leachate from living tissues) that the epiphytes are starved off the tree. In short, I suspect that it is a general example of the extreme case at Bako National Park, Sarawak; here, on a white-sand soil area, the only surviving epiphytes on upland trees were those associated with the nutrient-gathering ac- tivities of an ant colony (Janzen, 1974c). There are several conspicuous alterna- tive hypotheses as to why there is a shortage of vascular epiphyte biomass as compared to neotropical rain forest. a). The bromeliads ( Bromeliaceae) never made it to the Malaysian tropics and it is their absence that makes epiphyte biomass seem so low. Such a hypoth- esis does not explain why epiphytic orchids, gesneriads, ferns, asclepiads, Pipera- ceae, ericaceous shrubs, rubiaceous shrubs, etc. are equally low in biomass. The Dipterocarpaceae, which make up 30 to 90% of the crowns in the canopy of the forests I examined, have evolved bark traits inimical to epiphytes. It is certainly possible for this to occur, as there are species of neotropical rain forest trees that regularly have crowns clean of epiphytes while growing only a few feet from many species festooned with epiphytes. However, if this is the explanation, it is many more species of tree than just those in the Dipterocarpa- ceae that have perfected their anti-epiphyte defenses. It seems unlikely to me that a whole flora of large trees could evolve this ability. Furthermore, if nutri- ents are exceptionally scarce for epiphytes, then even weak defenses may be adequate to keep them of There are two 5 that are relevant to this hypothesis. On rare occa- sions I did encounter a native tree that was solidly covered with large epiphytes. For example, there is a medium-sized tree on the bank of the Tembeling River about halfway between Tembeling and Taman Negara that has hundreds of plants of a large basket fern on it (though perhaps it might be one huge clone linked by rhizomes). I saw no other individuals of this fern on the trees along the river. How does a huge plant like this one stay in the game as an epiphyte? It is possible that its litter-capturing leaves have an exceptionally robust ant colony living in them, or that its substrate tree is one of the few epiphyte-suscep- tible species in the region. Second, when Central American trees such as Pithe- cellobium saman (rain tree) or mahogany (Swietenia spp.) are planted in Ma- laysia, they develop large epiphyte loads. However, all the examples I saw were growing in small villages or roadsides where one would expect dust and other debris to provide high quality aerial fertilizer for the epiphytes. It is notable that these trees are deciduous forest trees in Central America and when trans- planted to the evergreen forest within their native country, they also develop exceptionally high epiphyte loads, even for the rain forest. In closing this section, let me call your attention to a quite different set of habitats where there appears to be a relationship between animal species rich- ness and harvestable productivity. On Costa Rican and Venezuelan mountain- side gradients, rather than the species richness of insects in sweep samples falling 714 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 off linearly with increasing elevation, it actually increases or at least stays about the same up to about 1,000-1,600 m elevation (Janzen, 1973a, Janzen et al., 1976). I have hypothesized that this “mid-elevational bulge” is the result of a similar bulge in harvestable productivity that occurs in the following manner. As one rises in elevation, the nights become cooler but the day does not cool as rapidly, and thus photosynthate production does not decline as rapidly as does nocturnal respiration. The result should be a greater amount of net produce per unit time for the plant, until an elevation is reached where diurnal photosyn- thesis is also severely reduced. If this is actually going on, should it result in an increase in numbers of species of herbivorous insects? I would argue yes, because as I have argued ear- lier, there should be more fractions of each plant species (e.g., the new shoot tips produced in the lower outer third of the crown) that are large enough to support a specialist herbivore. By like reasoning there should be an increase in the species richness of arthropod predators and parasites of these insects (Janzen, 1973a, 1977c). The more generalized a feeder (e.g., birds as contrasted with parasitic Hymenoptera), the less a taxonomic group should be affected, but all should be affected somewhat. Moving in the other direction, should the in- creased photosynthate production result in more species of plants than expected with a straight-line relationship between elevation and harvestable productivity? Yes, but again to a lesser degree than with the herbivores. By increasing, net photosynthate to an individual plant, there will be some kinds of specialization in which it can now participate (and thus more species can be packed into the habitat), but the relative heterogeneity of the increased photosynthate should be low compared to that received by the herbivore; the resource called sunlight is subdivided into many fewer compartments than the resource called "those plant parts that a herbivore eats." By these varied examples I mean to suggest that a very promising yet unex- plored area in tropical animal-plant interactions is the relationship between the pattern and amount of harvestable productivity and the numbers and kinds of animals present, and vice versa. We badly need solid data on relative abun- dances of animals and rates of production of harvestable parts of plants, col- lected with reference to particular questions such as “Given foraging inefficien- cies and temporal distribution requirements, how much small bird biomass can be supported by the understory fruits of Pasoh rain forest?" Or, “Do mid-eleva- tion plants replace shoot tips faster than their analogues at low elevations?” MONOCULTURE FORESTS In worrying about tropical forests, the intellectual interest of ecologists and population biologists has been largely focused on the “Oh My” habitats contain- ing many species of trees (e.g., Ricklefs, 1977; Janzen, 1970; Connell, 1971; Ashton, 1969; Grubb, 1977). However, what I find much more perplexing are the lowland tropical forests with extremely low species richness of large trees: Shorea albida peat swamp forests in Sarawak (Anderson, 1961, 1964); Mora excelsa forests in Trinidad (Beard, 1946; Rankin, 1977); Ocotea, Mora, and Eperua forests in Suriname (Richards, 1952); Gilbertiodendron dewevrei forests 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 715 in West Africa (Gérard, 1960); mangrove forests around the world (Watson, 1928); Strobilanthes forests in the Asian tropics and bamboo forests around the world (Janzen, 1976b); Raphia taedigera, Pterocarpus officinalis, Prioria copai- fera, and Parkinsonia aculeata swamp forests in Costa Rica (Janzen, 1977b and unpublished). For the mercenary at heart, esoteric ecological studies of these sites and their plant-animal interactions should tell a very great deal about the art of growing monocultures of tropical trees without a large input of pesticides, herbicides or other costs. Perhaps instead of worrying about the return to nature through the reinvention of mixed stands, which is so much in fashion these days, we should be studying much more intensely those species that naturally occur as pure stands. Not that I am eager to see this information become part of the foresters’ operating protocol, however, unless it is accompanied by a (unlikely) kick-back to biology in the form of inviolate forest preserves. The questions these pure stands bring to mind are numerous, and here I mention just a few. (a). From whence come the pollinators when a large pure stand suddenly comes into flower, a pure stand that has had little or no flowering activity for one to several years? In general, these monocultures are adjacent to much more mixed stands of plants, and I suspect that it is from these stands that they draw most of their pollinators (though bamboo use wind and Strobilanthes use highly nomadic bees; Janzen, 1976b). In the specific case of Dipterocarpaceae, which form a “monoculture” of sorts in Malaysian forests if the entire family is viewed as a species, Ashton and his associates have found that the enormous quantities of flowers suddenly produced are probably pollinated by thrips (Ashton, per- sonal communication), and I suspect that these insects feed on the vegetative parts of dipterocarps or other plants during the intervening years. Further, be- ing very small, thrips can have a very high rate of population growth; the flow- ering season for dipterocarps as a whole is 4 to 6 months in length and thus there can be an extensive population explosion of thrips. Finally, it appears that the flowering times of the different dipterocarp species are scattered through the overall flowering time (in contrast to the highly synchronous fruit drop over about 2 months ), and I suspect that this is the result of interspecific competition for pollinators (Janzen, 1977d). (b). In every monoculture stand of tropical forest known to me, the usual dispersal of seeds is by falling below the parent, variously aided by wind, thus a shortage of dispersal agents would not appear to be a problem for these plants. However, this kind of dispersal means that a near neighbor is likely to be a sib, mother, grandmother, etc., and that outcrossing is therefore more difficult than in a population whose seed shadows overlap widely due to animal dispersal of seeds. But then again, in a permanent monoculture, perhaps the best genotypes are extremely specialized to that site, and thus genotype disruption or offspring heterogeneity through outcrossing and/or interspecific introgression is more dis- advantageous than in a site that is more varied edaphically and more varied with respect to herbivore challenges. (c). Why don't those herbivores that can deal with the defenses of mono- culture tree stands move into these habitats and literally mow them to the 716 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ground? There is certainly no escape in space. There are herbivores that can feed on the vegetative and reproductive parts of these plants (e.g., Anderson, 1961). I suspect that the answers to these questions are in the following area. When the habitat first appeared, and various species of trees were specializing with respect to it, those that got mowed down when they occurred in pure stands probably dropped out of the race early on (they should still persist in mixed stands, however). Second, one of the traits for living in a habitat supporting a pure stand should be the evolution of those kinds of chemical and behavioral defenses so effective that the plant does not rely on escape in space; perhaps they are more expensive chemically, but then again perhaps they can be afforded owing to less investment in interspecific competitive ability (e.g., desert cacti). Third, these plants have escape in time, and they use it; in many species seed production is highly synchronized within the year and in many species, at supra- annual intervals (bamboo being the epitome; Janzen, 1976b). Owing to the extreme specificity displayed by many tropical seed-eating insects, the seed crop of the monoculture stand does not necessarily draw a guild of insect seed preda- tors in the same manner as the flower crop may draw a guild of flower visitors from the surrounding mixed forest. However, such plants may also be involved in satiation of seed predators, and then they may draw large numbers of animals from surrounding areas and require long supra-annual periods to accumulate enough reserves for enough seeds to satiate these animals (e.g., bamboo, Dip- terocarpaceae). The same process is likely to be operating with new leaf pro- duction. For example, in Corcovado National Park (Costa Rica), all trees of Mora oleifera drop their leaves in late November and put out a new synchro- nized crop in December. SECONDARY COMPOUND CHEMISTRY AND HERBIVORES This is undoubtedly the most actively expanding area in tropical (and extra- tropical) animal-plant interaction studies. It is easy to predict that the descrip- tive data and tests of hypotheses over the next ten years will make our current understanding seem amazingly primitive and naive. Take any paper on this subject in a current journal, and you can generate more questions with its data than it answers; and if not, just combine it with the next apparent test of the same hypothesis to get the desired effect. We are even still drowning in termi- nological difficulties, with specialist, generalist, secondary compound, herbivore, strategy, community, at the top of the list; at least we seem to have left niches by the wayside. It is presumptuous for me to finger promising areas, but since asked, I will presume. As I mentioned earlier, do not jump on me for leaving out yours, just be happy that it is not yet a bandwagon. I will simply ask questions, the answers to which I feel are either as yet invisible or are only dimly visible. (1). What sort of enzyme systems occur in the guts and livers of herbivores (e.g., Freeland & Janzen, 1974; Krieger et al., 1971), and can they be turned on and off so fast that an animal can move from one species of plant to another with hardly a pause? Yes (Brattsten et al., 1977). If they have them generally, 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 717 and can be so activated, why aren't all animals generalists that can feed on any kind of foliage? (2). Is it true that large animals mix their foliage intake as a way of mini- mizing damage from any one secondary compound (dilution, antagonisms, keep- ing each compound at a low concentration), or do they do it largely for nutrient balance reasons ( Westoby, 1974; Freeland & Janzen, 1974 (3). There is more foliage present of species that big herbivores are known to eat than they do eat; does this mean that they are not food-limited (Berwick, 1974), or does it mean that there are upper limits for even the acceptable items? Or does it mean that the critical times in food shortage (pregnancy, weaning, etc.) have not been examined? (4). How do the metabolic costs of making secondary compounds (Penning de Vries et al., 1974) compare with the fitness costs of making them? 5). At the habitat level, what is the structural array of secondary com- pounds? In other words, are defenses more heterogeneous within than between the food of herbivorous guilds (Janzen, 1973b; Feeny, 1976; Rhoades & Cates, 1976; Cates & Rhoades, 1977; Futuyma, 1976)? (6). Why do many plant parts contain trace amounts of a variety of second- ary compounds, and then a large amount of a few? Are they really sloppy or does this array present a more viable or effective defense against the more spe- cialized or the more generalized animals? 7). How would you design the optimal pathway for the production of a secondary compound? Example: minimize the number of places that enzymatic reactions are needed, maximize the number of times that the same enzyme can be used, canalize the substrate-product sequence such that its intermediate parts cannot be stolen from other pathways (but perhaps a really sophisticated system would allow borrowing? ). (8). Why are specialist animals specialized to only one kind of host (if they are); is it that detoxification systems are really that intersystem incompatible? Or is it like the canavanine system where the beetle is using that which is pro- duced by the detoxification process (Rosenthal et al., 1977), and therefore we can state that the animal avoids other hosts not only because they are toxic but because they do not offer special dietary input. (9). Why are sugars the molecules tacked onto large active molecules to render them inactive (e.g., lectins, Liener, 1976, and other glycosides)? Is it just because of the ubiquity of hydrolyzing enzymes in animal guts? (10). Since insects have a much shorter generation time than do plants, why don't insects always come up with a resistant strain that eliminates the plant before the plant can evolve a chemical defense? (11). What happens when we view cellulose as a secondary compound? (12). If bacteria can degrade cellulose, lignin and other such indigestibles, why can't animals do the same? — — Wuy po Fruits ROT AND SEEDS Mol? The brewery's fermentation vats are not where yeast evolved ethanol produc- 718 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 tion and bread is not the native habitat of blue bread mold. An exploration of the biology of rotting fruits and molding seeds is likely to produce some very interesting and unexpected results, both here and in the tropics. When a mi- crobe has found itself a ripe fruit, it has two options. It can begin using the resource rapidly and directly for its own growth and multiplication. On the other hand, it can do this somewhat more slowly and convert part of the re- sources into compounds which the anticipated vertebrate consumer of that ripe fruit will find objectionable, toxic, repulsive, etc. I suspect natural selection to have favored genotypes displaying the latter solution. The selection should be more intense the more rapidly the dispersal agents and fruit parasites remove the fruits from the tree, and the less fastidious are the frugivores. Seen in this light, a sour or alcohol-rich fruit is not just an accident of microbial metabolism or the detritus of microbe-microbe warfare, but may also be the explicit out- come of selection for avoidance of consumption of microbes (or insects) by vertebrates (Janzen, 1977f The same argument may easily be applied to the fungi whose hyphae grow over the surface of grain caches of man and other animals. Only here, they are protecting a much more valuable resource (higher nutrient content, lower 1585 dance, large amount of work invested in harvest is a measure of work that will have to be repeated to reharvest it if lost, reserves for a resource-poor future, etc.). The protection will have to be more violent than for a fruit, and it is; aflatoxins, ergot alkaloids, and antibiotics may be used as examples. It is of par- ticular interest here that both insects and vertebrates are susceptible to these compounds, and the only fungal hyphae that regularly make such nasty things are those that live on grain stores (Janzen, 1 The biology of rotting fruit and moldy grain stores in the tropics is unknown in the wild and in most human habitats. Why do cassava tubers (Manihot) spoil so quickly that a major portion of the earth’s cassava production land is serving as a storage bin because once harvested, cassava tubers have to be eaten? An- other way to ask this is "Why do the spoilage organisms in cassava tubers so quickly render them unuseable for vertebrates?" Many species of tropical fruits are notorious for being poor at shipping and storage, unless picked extraordi- narily green. Is this because the fruits are particularly susceptible to rotting organisms (owing to lack of selection for long half-life of ripe fruits owing to very active dispersal agent guilds), because tropical microbes are especially com- petent in their competition with vertebrates (owing to very active dispersal agent guilds), or a combination of the two? How long does a yeast clone have to make a tropical rain forest fig so sour that a bat will leave it on the tree? Perhaps it may have only one or two days at the outside. I should add that the opportuni- ties for coevolution of competitive partners against the vertebrates is very great, and may take the form of teams of microbes and the insects that carry them from rotting fruit to intact ripe fruit. It is even possible that an exploration of some of these esoteric areas might well lead to pragmatic applications in the area of Ashton's (1976) proposals for exploitation of wild tropical fruit trees for their fruits. 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 719 THERE Is No OPTIMAL SEED The fitness of the female parent tree has to be measured in some way rele- vant to the number of new members she contributes to later generations and how well these new members do on the same parameter. The environmental chal- lenges presented to her seedlings are varied and unpredictable with respect to any given seed (though the variation and relative abundance of environmental challenges may be quite predictable to the parent). There can therefore be no optimal seed size for any species of tree. A large seed may generate a very strong seedling, but it may be sorted out by the seed dispersal process so as to land in a poor site. A small seed may generate a puny seedling but regularly land in a heavily insolated site. This year, a female may have % of her seeds land on dry sites and % on wet sites; next year she may do the opposite purely because of interyear differences in weather. In view of these problems, I hypothesize that there can only be an optimal distribution of seed weights within a female's seed crop. If this is so, then the distribution of weights of seeds (or some other mea- sure of the size of the bag lunch for the seedling) is not a simple outcome of sibling competition and physiological sloppiness by the parent plant, but rather may also be engineered by the adaptive value of having various proportions of the seeds in different weight classes. Incidentally, since the challenge to seeds varies from year to year, there may not even be an optimal seed size distribution in a crop; it will have to be within a lifetime. Ateleia herbert-smithii, a caesalpinaceous legume tree in Santa Rosa National Park, Guanacaste Province, Costa Rica, may provide an example. Not only does this tree display 1.89-fold variation in the mean seed weights among trees, but within a crown there is 1.6- to 2.6-fold variation in seed weight. There are some cases where twins are produced in the normally single-seeded wind-dispersed fruits. Here, the pair of seeds has a combined seed weight greater than the mean weight of solitary seeds, but each twin weighs considerably less than the indi- vidual solitary seeds. This can be simply interpreted as the outcome of sibling competition. However, it could also be a mechanism for dropping some light seeds near the parent plant. A fruit with twins should fall in that part of the seed shadow that normally receives single-seeded fruits with heavy seeds. In fact, the production of twins may be adaptive simply in homogenizing the seed shadow (Janzen, 1977h). In the same vein, the 3.62-fold variation in seed weights found within a single crop of Mucuna andreana could well be adaptive in gen- erating a more homogeneous seed shadow than if all the seeds weighed the same (Janzen, 1977g). A small hard object moves through the digestive tract of an animal at a rate related to the objects volume, shape, specific gravity and (probably) surface texture ( Hoelzel, 1930; Alvarez & Freelander, 1921; Hinton et al., 1969). It fol- lows that if a vertebrate eats a distribution of seed sizes from a single crop, they are likely to come out in a different pattern than they went in. Further, the prob- ability that they will go in at all should be influenced by these traits, as well as the relative seediness of the fruit. For example, does a tapir spit out flat 500 mg seeds more frequently than spheroidal 500 mg seeds of the same species? Does a deer chewing its cud spit out the large members of a seed crop and let the 720 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 small ones pass on through? Could the small seeds from the same Enterolobium cyclocarpum seed crop crack between the molars of a peccary more easily than the large seeds? Yes (Janzen & Higgins, 1977 Once we put on this pair of glasses, interesting variation pops out all over the place. What is the meaning of variation in fresh ripe fruit weight within a trees crop? Bonaccorso (1975) noted that different species of bats took Ficus insipida fruits of different weights from the same tree; this should generate a quite different seed shadow than if all the figs were of the same weight and thereby taken by only one species of bat. What is the significance of the spatial variation in fruit location in the crown? What is the significance of the tempo- ral variation in fruit ripening times within the crown? What about the variation in seediness within a crown? But do not forget the zygote. The world is not constituted solely by female parental manipulation. What is good for the parent is not necessarily good for the individual seedling. Once the zygote is formed, you might argue that it should do everything it can to extract as many resources as possible from the parent plant. Of course, the parent has many physiological ways of controlling this, but there may also be good reasons for the offspring to constrain its glut- tony. For example, the very large seed may simply be spit out below the parent by the dispersal agent, and its less greedy sibs make a happy passage through the intestine to a distant light gap. The large seed may lower its fruit/seed ratio to where it is taken late if at all, and thus be killed by some seed predator taking what has been left behind by the dispersal agents. The individual zygote is not in the process of generating a seed shadow but rather in maximizing its own chances of survival to a highly reproductive adult; it has only one chance, the parent has many. — OPTIMAL MATE SELECTION Animals are conspicuous in their courtship displays, fickleness, promiscuity, coyness, variably intense rape, and other descriptors of mate selection by both sexes. What are the analogous processes in plants? I would like to suggest that they are choice of pollinators, timing of flower presentation, duration and time of stigmatic receptivity, duration and time of pollen release, degree of separation of the sexes within and between conspecifics, abortion of ovules, and abortion of zygotes of a variety of ages. The core questions are “How many of which fathers does the female part of the plant genome want for any given seed crop?” and “How many offspring in which and how many seed crops does the male part of the plant genome wish to sire?” I would argue that in either case the answer is not a maximum number but rather some optimal number and optimal distri- bution. Further, I see no reason to believe that the optimal numbers and distri- bution of fathers and mothers are very likely to be the same for the female and male, and that the difference leads to such things as differential pollen accep- tance, selective pollen presentation, monoecy, dioecy, etc. (Janzen, 1977a). I suspect that the most unappreciated mechanism for shaping the genetic composition of her seed crop is that of zygote abortion. It is conspicuous that the individuals of a very large number of species of plants regularly abort all but a 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 721 very small number of the flowers that they produce. Flower to mature fruit ratios of 100-500 to 1 are commonplace in large tropical trees. Even careful hand pollination does not drive this ratio downward, though it can in certain species. Incidentally, I should note that pre- and post-zygotic abortion are probably not as different as they would seem since both should be controlled by the seed- bearing plant. The male portion of the zygote is certainly not going to be selected for abortion tendencies as it has all to lose and nothing to gain. Since my focus is here on animal-plant interactions, let me list some of the ways they influence abortion of flowers and zygotes. (a). By attacking a fraction of the ovules and zygotes in the flower and developing fruit, seed predators may render the fruit not “worth” the cost of further development or maturation. By being unpredictable in which flowers or immature fruits they will attack, and to a certain degree in how many, such animals require the retention of a large number of (perhaps) suitable flowers or green fruits from which the fruits to be matured can be selected after the animals have taken their toll (the excess flowers or fruits are then aborted). c). By bringing variable amounts of appropriate pollen (e.g., pollen that is not from yourself or a close relative) to the stigmas, the pollinators require the production of a large number of stigmas so that at least some minimum number get the right amounts of the right pollen; the flowers containing stigmas with the wrong pollen are aborted. (d). Owing to the vagaries and resource-gathering behavior of visitors to flowers, a large flower crop may be necessary just to satiate visitors, to get the right kinds, and to do it in competition with other plants; these flowers are then aborted simply because insufficient resources can be spared to mature their seeds even if pollinated. Viewing pollen donation and capture in the light of these comments brings me to the problem of the act of not setting seed by a flowering tree. With dioe- cious species, the lack of seed production by male trees has never caused the puzzlement it is due (but see Bawa & Opler, 1975, 1977). Dioecy is just a more final example of the behavior displayed by a tree with perfect flowers that like- wise sets no seed. I would hypothesize that such trees, dioecious or hermaphro- ditic, have often made some kind of an internal decision that they will have a higher fitness by putting everything into pollen donation rather than into pollen capture and nursing zygotes. Medway (1972) recently commented on how “after flowering, invariably some species failed to produce fruit”; Grubb (1977) picked this up as “circumstantial evidence exists for massive failure of pollina- tion in some species . .. in the lowland tropics”; it is commonplace for foresters to label trees that flowered but did not seed as having “failed” to reproduce. Quite the contrary, they may have reproduced much more heavily than the tree bearing seed, simply by having sired many seeds on that tree and others that did set seed. We don't label an animal as having failed to reproduce because he doesn't get pregnant after copulating. There are at least two ways that animals may be responsible for morphologi- 799 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 cal or physiological dioecious behavior by plants, both of which deserve much more attention than they have received (a). If seed predators and their satiation are involved in the tree's biology, the tree reproducing by seed may have to produce a very large seed crop or none at all; here then, the highest fitness may be achieved by being solely a pollen donor when young, sick, or between large seed crops. (b). Dispersal agents may not be accurate enough to put more than an occasional seed into a habitat in which there are enough resources to be a healthy seed-bearer, but may place many seeds in habitats that just barely allow adult survival. The plant, once dispersed to the latter habitat, cannot get up and walk to a better place, and thus may be doomed to be a pollen donor or nothing. COMPLEX ANIMAL-PLANT ÍNTERACTIONS There are four complex animal-plant interactions about which we now know a fair amount: orchid-euglossine bees, figs-fig wasps, neotropical acacia-ants, and leaf-cutter ants. Their taxonomy is fairly well understood, the basic ele- ments of the interaction are understood, and they have been widely publicized. On the one hand, it appears that there is little interesting new ground to be plowed with each, and therefore the bright young field naturalist should look for other systems on which to expend energy. I would contend the opposite. These systems are now prime for high quality field studies incorporating mod- ern ecological and population biology thought; the student need not waste years doing their taxonomy and natural history just to determine where to start. I have not started such a new study with any of the first three, but by nibbling at their surfaces for just a moment, the following interesting areas appeared. The fourth is being heavily studied by several investigators and I will not dwell on it here (e.g, Rockwood, 1975, 1976; Cherrett, 1968; Martin, 1974; Lugo et al, 1973; Hubble, personal communication ORCHID-EUGLOSSINE BEE INTERACTIONS (a). How many parasites of the system exist (analogous, for example, to Pseudomyrmex nigropilosa in acacia-ants (Janzen, 1975a) or to Sycophagus sycomori in Ficus sycomorus (Galil et al., 1970)? It is assumed that the bees that come to the orchids and to the scents put out to survey them are pollinators of one or more orchids. There is no biological reason why this has to be so. Even if the bee does pick up pollinia, there is no guarantee that it has the appropriate behavior and/or morphology to put it back in the right place. It is already recognized that a visiting euglossine may not be the pollinator, yet get chemicals from the flower (Dressler, 19683; Dodson et al., 1969); however, there is no reason that such a bee has to be the pollinator of any orchid. However, there are obvious forces operating to set the carrying capacity of the habitat for such parasites, and this should vary with the number of species of orchids, the number of species of euglossines, their numerical relationships, seasonality, etc. (b). Is the visitation of an orchid's flowers, in its natural habitats, habitat- 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 723 independent? Some observers have already noted that certain orchid bees will not come to baits or orchids placed in the open sun but will visit them in the shady nearby rain forest understory. I suspect that the story is much more com- plex than that, and much more interesting. In March 1977 I put five different chemicals (cineole, eugenol, methyl salicylate, benzyl acetate, and methyl cin- namate) out in five different forest types on the same day within a circle of 2 km radius in Corcovado National Park, Costa Rica. The numbers of euglossines that came to each site differed dramatically and the array of species differed somewhat, suggesting that if any given orchid were growing in one of these sites, its visitors could have been dramatically different. An orchid is not an orchid is not an orchid. Such differences should have dramatic effects on species packing in orchids, inter-habitat species richness of orchids, and the relative fitness of an orchid in a given habitat as compared with a conspecific in another nearby habitat. (c). How much self-pollination occurs in hermaphroditic euglossine-polli- nated orchids by the simple event of a bee picking up a pollinarium today and bringing it back tomorrow? At least one tropical orchid, Encyclia cordigera, is highly self-compatible (study in progress, Santa Rosa National Park, Guana- aste Province, Costa Rica). Since most adjacent conspecifics are probably closely related (orchid seed shadows are probably strongly peaked as in other wind-dispersed seeds), there is also likely to be an extraordinary amount of incest in orchid matings. And if interspecific hybridization occurs physiologically so easily with or- chids (Dressler, 1968b), does this really mean that they have extremely faithful pollinators as is generally assumed, or does it mean that orchids live in a world where it is very profitable to steal genetic information as whole blocks or as proven mutations from other genomes? If they do the latter, then I expect strong selection for the ability to obtain this information without severe perturbation of the phenotype. The conventional means for detecting hybrids, the means that are used to say that orchid hybrids are very rare in nature, might be therefore of little or no use. d). An orchid female generally has fewer fathers for her clutch than does a member of any other family except perhaps the Asclepiadaceae (and speaking of which, what do orchids and asclepiads ecologically have in common so as to have generated their convergence on this axis?). Every orchid fruit, with its hundreds of thousands of seeds, has but one father. Depending on the behavior of the orchid bees and the density of orchids in the area, even if there is more than one pod per plant, it is possible for multi-podded clutches to have only one father. At the very most, the ratio of fathers to seeds has to be on the order of a very few to hundreds of thousands. In what way could this observation be related to the point made in the last paragraph of the previous section: FIGS-FIG WASPS (a). Since no one has ever done anything but rear the fig wasps out of syconia (and none of that has been done in a quantity any greater than that needed for taxonomic purposes) (Hill, 1967; Ramirez, 1970a), there is no infor- 724 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 mation on what kind of interspecific pollen is being carried into a fig crop. Again, the literature loves the statement that hybrids of figs are very rare (e.g., Hill, 1967); however, since there is only one experimental cross on record ( Con- dit, 1950) how is one to know what a hybrid fig looks like? An extensive exam- ination of the remains of the female fig wasps inside a large number of recently pollinated but still quite immature syconia would tell how much foreign pollen is getting into the system at each generation. (b). Female fig trees pay offspring for pollination. Fig wasps are seed pred- ators. There is no published study of the intensity of this predation, but one in progress shows 30 to 50% for three fig species. What is the possibility that one of the selective pressures favoring gynodioecious figs (one morph of the popu- lation has syconia with solely female florets with styles too long for oviposition) is the act of obtaining the services of the wasp without paying the cost in zy- gotes? What is the overall cost in zygotes per intact seed for monoecious as compared with gynodioecious fig species? (c). Who eats figs? Everybody does. Nonsense. Yes, there are fig species that produce large numbers of small figs that are taken by a very large disperser coterie. These species must have very homogeneous seed shadows (generated by birds, bats, pigs, primates, etc.). On the other hand, there are fig species that seem to be largely visited by a very select subset of the frugivores in the com- munity (e.g. Ficus ruginervia produces large figs apparently taken only by gibbons, siamangs, and two species of squirrels when growing only a few meters away from Ficus sumatrana which was visited by at least 25 species of birds and 9 species of diurnal mammals—to say nothing of bats, which were apparently unrecorded; McClure, 1966). Morrison (1975) noted that the ripe figs of sev- eral species of Barro Colorado Island figs were taken largely or exclusively by bats; the howler monkeys are the only other visitors mentioned, and they ap- peared to take largely immature syconia. It is my guess that they took almost entirely immature syconia. I suspect that the traits of ripe fruits are engineered by the need to keep them out of the wrong dispersal agents as much as to get them into the right ones (and see Howe, 1977). There should be many parasites present in any disperser- fruit system. The trick is recognizing the parasites (as separate from the seed predators) because they do their damage by putting the seed in the wrong place, rather than by killing it directly. The neotropical oil birds (Snow, 1962) come to mind as the most glaring example, since it appears that many of the seeds they eat are later regurgitated in a cave where they die. Likewise, a Mexican oriole that eats the fruit pulp around Acacia cornigera seeds and drops the seeds below the parent tree may have killed those seeds as dead as if it had ground them in its gizzard. It appears that Andira inermis fruit pulp may contain an antibiotic which thereby renders it a high quality food item solely for those ani- mals that depend little on bacterial degradation for the extraction of nutrients from their food (Janzen, 1977e). It occurs to me that the various bat-dispersed figs on Barro Colorado Island may be doing the same; interestingly, such a com- pound might also be functional in slowing the rate of spoilage of a fig that has already been opened to the outside world by the exit of the fig wasps. If this 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 725 hypothesis is correct, then consumption of green figs before such a compound is activated might be the only way that a howler monkey can eat them, since it depends heavily on its bacterial community for degradation of foliage. Howler monkeys often avoid ripe fruits but eat the same species when green (Glander, 1975a, 1975b, 1977). d). The current consensus seems to be that fig wasps are very host-specific and that the figs are strongly synchronized within a given crown (Ramirez, 1970b; Hill, 1967). I have no doubts that this is the case for many species in many habitats. However, Ramirez (1970b) has already pointed out that on islands there may be selection for the loss of synchrony within a crown if the fig population is so small that there could be times when no tree is in a receptive state. There should be other habitats where both the host-specificity and the synchrony should break down. Butcher (1964) stated that the two native species of Ficus in Florida, F. aurea and F. laevigata, are pollinated by the same species of wasp, Secundeisenia mexicana. If this should occur anywhere, it should be Florida with its killing frosts. McClure (1966) noted that in the dipterocarp forests around Ulu Gombak there was a two month progression of ripe figs through the crown of a Ficus sumatrana (and shows it in the graphs for two other species as well). Fig wasps take about a month for a generation. Unless the pollination is synchronized but the syconial ripening grossly asynchronous, there is a very great possibility of self-pollination in these species of figs. At least two selective pressures could be operating to produce asynchrony within the crown. First, it may be that fig tree density is generally low in this forest, perhaps even as low as on a small island from the viewpoint of the wasps. Sec- ond, it may be that the actual biomass of fig-removing animals is small, and if all the figs this tree can make in a pulse were to be matured in a week or two, most would rot on the tree or the ground for want of not having been removed (the frugivores having been satiated). ACACIA-ANTS (a). There is no wild plant species anywhere in the tropics for which even an approximate herbivore load has been described. If the ants are removed from neotropical swollen-thorn acacias, the herbivores that normally feed on them and those that feed only on the largely undefended plants often become tem- porarily abundant and easy to census. Combined with careful observations of the insects feeding on acacias with their colony intact, it would be possible to not only rapidly identify the herbivore load of at least one plant species, but to ask how it behaves when the defenses of the plant are suddenly removed with- out physiologically altering the plant. When my early ant-acacia studies came out, there was a good deal of *Oh My" of how all those little ants could do such a marvelous job of defending the tree. The significance of those studies lay not, it seems to me, in this aspect. Rather, here we have a plant that in nature can be deprived of its defenses and thereby demonstrate how important are its de- fenses in determining the amount and structure of the herbivore load. (b). When an ant-acacia is crossed with a non-ant-acacia, the offspring are most amazing organisms. They have either Beltian bodies, large to normal 726 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 — thorns, large nectaries and seem to be poorly protected, or they almost entirely resemble the non-ant-acacia parent. Thus it is as though the ant-acacia traits were one gene (and you get an ant colony as well). If this gene is donated through interspecific introgressive hybridization, a non-ant-acacia may become an ant-acacia. It does this without ever losing its other traits which, for example, would cause it to be placed in a quite different subgroup of the genus. It would then appear that the ant-acacia interaction had evolved independently on sev- eral different occasions. The 10 or so species of neotropical ant-acacias seem, on the basis of flower and fruit traits, to belong to quite different species groups (Janzen, 1974d). The system is wide open for experimental verification. And what would it mean to attempts to reconstruct phylogenies based on suites of traits? (c). There are three species of obligate acacia-ants in Guanacaste Province, Costa Rica, that protect the common swollen-thorn acacia, Acacia collinsii (Pseu- domyrmex belti—black; P. nigrocincta—the smallest of the three and with a yellow body with the gaster held straight out in back; P. ferruginea—rust red with the gaster commonly curled under the posterior part of the thorax). They all eat the same thing (Beltian bodies and foliar nectar), live in the swollen thorns, and aggressively protect the plant. All three occur in every habitat I have ever sampled. No large colony will cooccur in the same acacia with another colony. All colonies have but one queen, and while many queens may start out in the same acacia seedling, the winner takes all. It is hard to imagine a more monomorphic resource than ant-acacias. I suspect that originally one species was a specialist on the ant-acacias in the forest understory (P. nigrocincta), one spe- cies was a specialist acacia growing in open but moist sites such as river edges and marsh edges (P. belti), and one species was a specialist on acacias growing in fully insolated but very dry sites (P. ferruginea). When the pasturing, crop- ping, and timbering broke up the habitat structure, all three species moved into each other's habitats and are found there today. In contemporary habitats the ratios of the species vary widely among habitats, and I suspect that in the origi- nal habitats there were always a few colonies of the other two along with the most abundant species. The colonies can be moved about, the acacias can be secded with queens, and all occur in large numbers. The opportunities for experimental study of direct competition between sessile animals in terrestrial community are enormous. ). The pulp around the seeds of Acacia collinsii, moist when ripe, can be 1 in a plastic bag, sealed, and left for a year without spoiling. From what I said earlier about bats and Andira and fig fruits, there is one obvious sugges- tion about what it contains. All swollen-thorn acacias have seeds imbedded in a sweet pulp (Janzen, 1974d), apparently for seed dispersal by birds (in contrast, I know of no neotropical non-ant-acacia with this trait). Presumably this trait arose many times independently. What a marvelous opportunity to study con- vergence in fruit protection traits. The above hypotheses and systems briefly alluded to for orchid-euglossine bees, figs-fig wasps, and ant-acacias are, I am certain, only a tiny fraction of the studies that can be developed into large and clean studies in coevolution, popula- 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 727 tion biology, gene flow, competition, etc. These suggestions are possible because some background has now been developed for these systems. Yes, the “Oh My” part of the studies has been killed rather dead; now it’s time to start on the interesting things. ONE LINERS (1). Why do trees have rotten cores? Hypothesis: to provide a place, through the removal of unneeded structure, for animals to roost and defecate and for microbes to grow, which in turn generate a nutrient pool for the tree's roots. The actual process should be through the selective and only temporal protection of the core of the heartwood from decomposers (Janzen, 1976c; Fisher, 1976; Thompson, 1977). ). Why do vertebrate dispersal agents leaves the tree to eat the fruit they have picked? Hypothesis: because there has been strong selection for chemical, morphological, and behavioral traits of the parent tree to be an objectionable place to perch. The competing hypothesis is that the fruiting tree is a focal point for foraging carnivorous predators. 3). Why don't the ants of the world take over the flowers of the world and protect their nectaries just as they do extra-floral nectaries (even on those on the outside base of the flowers)? Hypothesis: there is an anti-ant compound generally present in floral nectar. And in case you still think nectar is just sugar water, read recent papers started off by the Bakers (Baker & Baker, 1975; Baker, 197: (4). Why do rain forest seedlings with mycorrhizae recover from herbivory much better than do conspecific seedlings that have not yet acquired (accepted?) a mycorrhizal association? Hypothesis: the seedling can mark time waiting for the appropriate fungal associate using only its seed reserves plus the very small amount of resources it can harvest in the heavily shaded rain forest understory, but if it has to undergo the major capital investment of replacing lost photosyn- thetic structures, it does not have adequate resources (Janos, Ms (5). Why do rain forest understory shrubs contain a high amount and diver- sity of so-called trace elements (boron, cobalt, etc.; F. Golley, personal commu- nication). Hypothesis: the heavily shaded rain forest understory is one of those resource-poor habitats where chemical defenses are of utmost importance (Jan- zen, 1974b); large quantities and many kinds of secondary compounds may re- quire large quantities and many kinds of co-enzymes for their protection, and co-enzymes normally contain a molecule of a so-called trace element. (6). What is the distribution of intensity of seed mortality by animals among the members of a tropical tree population? Hypothesis: there is a very skewed distribution, with most individuals producing few or no surviving seeds, and a very few producing most of the members of the next generation. If this is veri- fied, then the opportunities for rapid genetic change and the selection for high levels of information exchange among members of the population should be very high. (7). What do whole disperser coteries, herbivore loads, and suites of polli- nators for an individual and a population look like? Hypothesis: they will be 728 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 rich in species that parasitize the system, rich in species that gain resources but take so little that the mechanisms to remove them would cost much more than the value of what they take, have a few key species that drive many of the traits of the system, treat individuals very differently even within populations of closely adjacent individuals, and display strong competitive interactions through the medium of the plant (Janzen, 1973c), as well as directly with each other. (8). Why are grasses so edible? Hypothesis: they are involved in satiation of leaf predators (being possible by having put the high investment centers underground where they cannot easily be reached by fire and herbivores); the cost of chemical defenses would generally be higher than the fitness loss that occurs through the grazing that is produced by the animals that make it from one rainy season to the next (and see especially Sinclair, 1977). (9). Are plant apparency arguments (Feeny, 1976; Rhoades & Cates, 1976) applicable to all parts of plants, as they appear to be to foliage? Hypothesis: probably, but it will require some very careful definitions of what is apparent and what is not. For example, at least two major groups of mast-seeding trees, oaks and dipterocarps, have polyphenolics as their major chemical defenses (if any be). A very large number of trees that do not display habitat-wide supra- annual synchrony of seed production have alkaloids and other conventional tox- ins in their seeds. One could argue that the seeds of mast-seeding species, widely spaced in time, are less apparent than the seeds of tree species that fruit or seed every year. However, I could rebut this argument by noting that when the mast- seeders do seed, they do it in such abundance (and often in such pure stands ) that the seeds are enormously apparent; on the other hand, species of seeds defended by alkaloids, etc. are often very widely scattered or rare, and thus much less spatially apparent, even if they fruit every year. In closing this sec- tion, I must note, however, that many species of seeds contain both direct toxins (alkaloids, uncommon amino acids, cyanogenic glycosides, etc.) and digestion inhibitors (tannins, lectins, protease inhibitors); e.g., chocolate beans contain tannins and 3 kinds of alkaloids. Perhaps it is that as the nutrient content per bite of food rises, the adequacy of only one class of defense in the plant part declines precipitously. 10). Why is the biomass of palms in Africa so low? The exceptions are oil palm (Elais guianensis) and raphia palm (Raphia taedigera) pure stands in swamps, borassus palms in very arid areas (e.g., Samburu National Park, Kenya), and very thorny climbing palms in swampy rain forest. Richards (1973) has already noted that Africa has a ridiculously low number of species of palms— about 50 species as compared with about 1,140 in the neotropics and 1,150 in the Asian and Australasian area. Hypothesis: palms, with their single large growing points, are particularly susceptible to herbivory by elephants, mammals which were until recently prominent browsers in African forest habitats. (11). Can the fitness of a plant be raised by herbivory, thereby selecting directly for palatability or lack of defenses in a plant part? This hypothesis has recently been championed by Hendry et al. (1976), Owen & Wiegert (1976), and Harris (1973), among others. Aside from the obvious cases of seed dispersal and pollination systems, and the problems of defenses in these systems being incom- 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 729 patible with “allowing” herbivory by the “appropriate” animals, I have yet to see a convincing case of a positive answer to this question. Of course, an herbi- vore may remove a part that would otherwise have to be actively dehisced by the plant, or turned off by the plant, but to put such an activity in the hands of the herbivore requires that it will only do that and that it will be reliable in its activity; that is to say, the plant loses part of the control over itself. In the most dramatic case, mild defoliation of crop plants may result in overall increased yield per field (Harris, 1973). However, this is quite easily explained by assum- ing that the defoliation breaks apical dominance, something that would result in loss of status in the natural competitive situation, but is optimal for properly spaced plants in the field situation. The same applies to cases where mild brows- ing of bushes appears to raise their vegetative productivity, and when mild defo- liation of a wild plant increases its seed production (e.g., Cavers, 1973). In short, just because a person runs faster after hitting a wasp nest, we do not conclude that (a) being stung raises your fitness and (b) susceptibility to being stung is a mechanism evolved by humans to get themselves to run. Finally, I can simply state that the various schemes frequently proposed for the “value” of herbivores to the ecosystem at recycling leaf contents are evolutionary nonsense. There is no evidence that a plant gains more from having its leaves eaten and then (perhaps) taking up some of the mineral contents of that leaf or feces from the litter below than from keeping its leaf intact in the first place. (12). Are extant gymnosperms and other “primitive” plants really freer of herbivores than are angiosperms (Regal, 1977)? If so, is this due to their sec- ondary compound chemistry or is it due to accidents of host location by herbi- vores? Are we seeing the bare remnants of a once great flora being pushed out by the combined actions of herbivores and competition from plants with a supe- rior growth form? (13). Prominent and severe defoliation by highly host-specific Lepidoptera and Coleoptera occurs during the first 1-2 months after the rainy season begins in the Costa Rican lowland tropics (see Rockwood, 1973, for examples of its effects). Hypothesis: the cessation of defoliation after a single generation of a given species of herbivore is due to the accumulation of secondary compounds (such as digestion inhibitors, see Feeny, 1976) in maturing leaves which makes them unavailable to the insects. The corollary of this would be that a tree is susceptible to this event only because it has to make a new crop of leaves each year. Opposing hypothesis: the cessation of defoliation after a single generation of herbivores is due to a buildup of parasites and predators at this time of year. There is no direct or circumstantial evidence to support this hypothesis. ew leaf-cutter ant fungus gardens are established by vegetative propagation from cuttings carried by the newly mated queens. Hypothesis: over many generations of leaf-cutter ants in an area, all the colonies will eventually be owned by the same subdivided individual fungus, which should in turn be the genotype that does best on that particular mix of plants which are available to the ants. This then becomes the world’s largest fungus and uses armies of leaf-cutter ant colonies to feed itself; will this result in manipulation of the colo- nies at a density and degree of intercolony aggressiveness which is suboptimal 730 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 for the ants and optimal for the fungus? Does this mean that leaf-cutter ants are more finely tuned to a habitat than expected, and therefore will also have more than the expected difficulty in colonizing new habitats? It appears that the species richness of herbivore loads of perennial plants reaches an asymptote within a few hundred years after the species has been introduced, with the level set largely by the areal extent of the population (Strong et al., 1977). Hypothesis: this conclusion will be most robust with highly apparent plants that are generally defended by digestion-inhibiting chem- icals, since it is these plants that will have the most in common with those native plants that are fed on by generalists; newly introduced herbaceous and other plants defended largely by more direct toxins should require considerably more time to accumulate a normal herbivore load from the indigenous pool, as this will require evolution on the part of local herbivores (Gilbert, 1977). Herba- ceous crop plants, however, will not be a useful test of this hypothesis; they have had their defenses bred out of them to various degrees and thus should quickly accumulate their saturation herbivore load. I should also note that a newly introduced wild plant will not likely occur in a major monoculture, as is the case with crop plants. Therefore, requirements of herbivore coevolution with the hosts physiological behavior, size, phenology, etc. may become much more important in slowing the rate of accumulation of the herbivore load than Strong et al. (1977) found to be the case with crop plants. (16). On 11 August 1977 a healthy male tapir swallowed 95 intact seeds of Enterolobium cyclocarpum and 80 intact seeds of Cassia grandis (weight about 0.7 and 0.5 g each, respectively); for the following six days there was no trace of these extremely hard seeds in the tapir's feces except for two E. cyclocarpum seed coats. Hypothesis: the seeds were sufficiently slowed in their passage through the tapir's digestive tract such that they were sufficiently softened such that they were digested rather than dispersed. The various eddy currents ( e.g., loops in the intestine) and pockets (e.g., caecum) could thus be highly adaptive in aiding digestion of seeds too hard to break with the teeth. Furthermore, large animals commonly thought to disperse hard legume seeds may well be extract- ing a high price in seed predation; there have never been studies of what per- centage of the seeds ingested actually survive the voyage through the animal. IN CLosinc I cannot resist commenting on the classes of administrative effort that I feel we lack in tropical animal-plant studies. First, I feel that we have quite enough hypothetical biology on the books. We desperately need information on the pragmatics of what is actually happening out there. I can generate, with a little help from my friends, a computer model that will predict anything; for example, a model can predict that increased productivity should increase species richness and that increased productivity should decrease species richness, or it can pre- dict that a predator should increase its specificity as prey gets scarce and that a predator should decrease its specificity as prey gets scarce. It all depends on what natural history facts you plug into the assumptions. Let's go out and get those facts, and ask what is their frequency distribution among real mem- 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 731 bers of real habitats and guilds. Of course, we should be gathering the facts in respect to questions, but let's not let the airy-fairy castles in the sky block out the sun. Second, many kinds of tropical animal-plant studies involve systems with patterns or cycles that will not be apparent until tens of years of data on the same individuals in the same habitats by the same investigators have accumu- lated. Funding for more than three years, and often more than two, is largely nonexistent unless you pay for it out of your own pocket. NSF states that they cannot tie up funds for long periods. Well, if NSF will give me $40,000 in direct costs to spend this year, there is no reason why it should not give me $40,000 in direct costs to spend over the next ten years in $4,000 per year bits. They have paid out the money the first year and that is that. I would like to explicitly appeal for the establishment of grants of that structure, grants that will float with the investigator wherever or whenever transient. There are many long- term studies that I am now setting up for the last 30 years of my research life that could have had another 12 years on them had this sort of funding been available for this explicit purpose in 1965. Without this funding, we have the ironic situation that the shorter the time I have left to do research, the more likely my funding is to be sufficiently secure that I can set up such studies with- out having to worry about having the funds to census them annually. Third, I would like to repeat a call (Janzen, 1977e) for some kind of inter- nationalized and centralized chemical identification service, analogous to the great museums and their contained identification services. Secondary com- pounds and nutrient analyses of plants are to animal-plant interactions what Latin binomials are to ecology and evolutionary biology. A contemporary tra- dition is developing whereby natural products chemists are being prevailed upon with ever-increasing frequency to do secondary compound determinations by ecologists and evolutionary biologists. And in a manner exactly analogous to whole-organism taxonomists, the natural-products chemists are being swamped. With a very few exceptions, and these exceptions tend to have a very short half- life for obvious reasons, their work is slow, interrupted, variable in quality, and heterogeneous in coverage. Their primary commitment is not to those field biol- ogists who send in a box of this or that at highly unpredictable intervals. Vir- tually all identifications are done gratis as a personal favor. When there is more than one class of secondary compound in the plant part, and this is nor- mally the case, the specialist concerned can readily isolate and identify within only one class. It is as though I had sent a tanager gut off to a museum and the determinations came back reading 15 Solenopsis geminata subsp. goofus, 12 creepy-crawlies, 14 slimies, 1 blob, and 102 hardies. There is no Museum of Secondary Compounds nor is there any laboratory in the world that for a rou- tine service charge will survey plant samples for kind and concentration of secondary compounds. Yet if entomologists, ornithologists, primatologists, ecologists, etc. are to give secondary compounds the attention that they have long deserved in understand- ing animal-plant interactions, such an identification service is essential. I am certain that the highly inconclusive nature of the tens of thousands of pages of 739 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 data that have been gathered on feeding biology of herbivores is largely due to (1) the impossibility for humans to identify secondary compounds with their own senses, and (2) the difficulty of getting such compounds identified by other workers. I have seen much evidence that field workers would be quite willing to conduct the feeding experiments and observations to place secondary com- pounds in their proper perspective if they could get them identified easily. If connected with the appropriate institution, I suspect that such a labora- tory could be established for less than 1 million dollars, and I suspect that its running costs could be largely met through charges for determinations. A spin- off would be research on the new compounds encountered and the purification (at cost) of large amounts of certain compounds to then be used in field trials. Such a facility won't happen unless some small and dedicated body of people take it on (e.g., people centered around E. A. Bell, Kings College, London; T. Mabry, University of Texas, Austin; P. Waterman, University of Strathclyde, Glasgow; R. Cates, University of New Mexico), and they won't take it on unless ecologists and evolutionary biologists can create a climate in the funding agen- cies for its support. So in conclusion then, where are the promising new areas in tropical animal- plant biology? Take systems that are already very well known in terms of general natural history and taxonomy, and apply current concepts of ecology and popu- lation biology to them, rather than picking on one of the many largely unex- plored “Oh My” systems. (2). Figure out how tropical plants survive in pure stands, rather than Worry about the mixed species stands. Use the organisms to tell you about the rates and kinds of harvestable productivity, and work backwards from this to expose the underlying causes; again, for the mercenary at heart there may be some powerful lessons here on how to competitively exclude our competitors or increase the yield from appar- ently low productivity sites (e.g., a rubber plantation may be such an example, discovered quite accidentally). (4). Apply the multitude of hypotheses and ideas that are appearing in secondary compound chemistry to tropical plants and the animals that feed on them. Do not be too fascinated with the generalities; let's get some frequency distributions of results first. Furthermore, it won't happen unless we can get some sort of an International Museum of Secondary Compounds, Isolation and Identification. (5). Stop thinking in terms of optimal seeds, fruits, flowers, reproduction times, seed crop sizes, etc. We have to start thinking in terms of optimal distri- butions in space and time for these parameters (as well as for other parts of plants and animals). Most plant parts are confronted by a set of animalian challenges or mutualists, not just one. (6). At the risk of being labeled a Darwinist fanatic, I would emphasize the value of looking very hard for the adaptive significance of traits that we regularly take for granted (e.g., variation in seed size within a seed crop, rotting of fruits, duration of ripening times for fruits, seediness of fruits). It has been 1977] JANZEN—ANIMAL-PLANT INTERACTIONS 733 my general experience that pessimism about the adaptive significance of a trait is strongly correlated with ignorance of the natural history of the organism. l the contemporarily fashionable ideas about parental investment, optimal parentage, sibling rivalry, etc. all apply to plants as well as to animals. In the tropics, animals play an enormous role in plant breeding systems and much of their obscure interaction with plants may become clearer when we come to understand what are the driving forces that determine which plant is to mate with which plant. (8). We need much, much more natural history of tropical plants and how they interact with animals. I don’t mean miscellaneous field notes of which beetle was found sitting on which plant, but rather natural history directed at interesting questions in ecology and evolutionary biology; we have plenty of them. LITERATURE CITED ALVAREZ, W. C. & B. L. q ps, 1921. 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Platt (1974) and Jordan (1975) have argued that this definition is inadequate, because a large portion of all biological and agricultural research and a significant fraction of medical and engineering studies can be construed to be studies of the interaction between organism and environment, whether or not the studies are really ecological. For example, a sewage engineer might convince a local town council that he is an ecologist because he studies interaction between bacterial concentrations and river flow, and therefore he could claim competence to prepare an environmental impact statement on the effect of sewage outfalls from new housing developments on the stream that passes through the town. We would argue that he is not competent because ecological problems resulting from sewage disposal are not limited to bacterial concentrations, but include such phenomena as eutrophication and resulting changes in fish populations, and recreational and economic use of the unpolluted river. If “the study of interactions between organisms and environment” is an inadequate definition of ecology, because many diverse types of scientists study such interactions, what then is the unit of study that is unique or basic to ecology? One system of classifying units of biology is the hierarchical approach. In this system, for example, the basic unit of study for cytologists is the cell, and the basic unit of study for the morphologist is the organ. For ecologists, the basic but it must have definable limits inside of which there are integrated functions. ecosystems.” If our definition of ecology is “the study of ecosystems,” we must then define ecosystems. An ecosystem is an integrated unit, consisting of interacting plants and animals whose survival depends upon the maintenance of biotic and abiotic structures and functions. The unit does not necessarily have to be isolated, but it must have definable limits inside of which there are integrated functions. What are these ecosystem functions? There are three functions upon which ecosystem ecologists focus their at- tention: energy flow; nutrient cycling; and water flux. Nutrients, energy, and water also are studied by physiologists, but what sets ecosystem ecology apart is the structure that supports these functions. Physiologists study flows of energy, nutrients, and water in individual organisms, whereas ecologists study them on an ecosystem scale. In using this definition of ecology we do not mean to say that the only truly ecological studies are those which follow energy, nutrients, and water through 1Centro de Ecologia, Instituto Venezolano de Investigaciones Cientificas, Apartado 1827, Caracas, Venezuela. ANN. Missouni Bor. Garp. 64: 737-745. 1977. 738 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 ecosystems. We certainly do not exclude those scientists who focus on species interactions and population dynamics. What is important is that the investigator maintains a holistic perspective. Maintaining a holistic or ecosystem perspective, in the sense that the scientist considers all ecosystem aspects, is what sets an ecologist apart from scientists of other disciplines. For example, if we see a scientist studying the aquatic life in a river close to a sewage outfall, we could tell if he has the ecological per- spective by asking him his objectives. If he answers that he is trying to protect human health by getting rid of harmful bacteria, he may be an environmental scientist, but he is not an ecologist because he is not considering all the ecological effects of the sewage entering the river. If he answers, in effect, that he is trying to protect human health by keeping man’s life support systems functioning, in this case perhaps by preventing eutrophication and thereby maintaining a down- stream fishing industry, he has the ecological perspective. He considers the implications of the problem, above and beyond the direct and obvious problem. A TnoricAL ECOLOGICAL HYPOTHESIS Because the title of this symposium is “Perspectives in Tropical Botany,” we should relate tropical botany to ecosystem research in the tropics. Tropical plants are, of course, the base of the food chain in tropical ecosystems. Tropical plants also recycle nutrients from decomposing organic matter on the forest floor and make the nutrients available to the animals. It is in this sense that we tie tropical botany to tropical ecology. We are going to talk mainly about our own ecosystem project in the tropics, but we will place it in perspective by comparing it with other tropical ecosystem research projects. The overall objective of our project is to study the structure and function of an Amazonian rain forest, so that increased ecological under- standing of the ecosystem can contribute to more effective applied management in future years. However, we are not interested only in the applied aspects, we are interested in the basic nature of Amazonian ecosystems, what temperate-zone man expected to find there, what he actually found, and how the differences can be explained in terms of ecosystems theory. Temperate-zone man has equated tall forests and large trees, and diverse flora and fauna with productive landscapes. When he encountered the Amazo- nian rain forest, he was impressed by the mass of vegetation and variety of orga- nisms, both of which exceeded his temperate experience. He concluded that the tropics must be very productive. However, when he converted tropical forests to agricultural plantations, yield declined drastically. Why? Temperate experience suggested that the yield was related to soil fertility. Therefore, the problem must be in tropical soils. And indeed, the amounts of essential nutrients could be very low. But then, how could luxurious tropical forests survive on such poor soils? Ecologists have hypothesized that development and survival of lowland tropical rain forest is through nutrient-conserving mechanisms that maintain the essential elements within the biomass of undisturbed forests, and that the destruction of these mechanisms by cut-and-burn agriculture results in rapid 1977] JORDAN & MEDINA—ECOSYSTEM RESEARCH 739 loss of nutrients, with a resultant loss in ecosystem productivity. While this con- cept is almost popular knowledge, the hypothesis has never been tested. The major emphasis of our Amazonian ecosystem research program is to test this hypothesis and to identify the nutrient conserving mechanisms which operate in the undisturbed tropical forest ecosystem. THE SAN CARLOS PROJECT The field site of our project is near San Carlos de Rio Negro, in Amazonas Territory of Venezuela. The site is within the north-central drainage basin of the Amazon River. There are two principal forest types in the area, both about equally important in terms of area. One is the tierra firma forest, located on laterite covered with a thin layer of sand or gravel. Species diversity is high, and biomass is close to 400 t/ha (Jordan & Uhl, in preparation). The other type is located on sand, with a podsol B horizon at about one meter depth. During heavy rains, the water table reaches the soil surface in this type. Biomass and species diversity is less than on the tierra firma site (Klinge, 1976). The experimental approach is as follows: We have a series of experimental and control plots on both soil types. We have measured the nutrient inputs, outputs, storages, and transfers of the major ecosystem compartments in these plots for one year. After one year, the experimental plots were cut and burned following the traditional local practices. In the podsol site, some areas were planted to rubber plantation and others were abandoned for secondary succession studies. In the tierra firma site, the experimental area was planted with typical crops of the area, manioc, pineapple, plantain, and a few other species. As a result of our observations and measurements during the first three years of the project, it has become apparent that a well-developed root mat and humus layer which occurs on top of the soil surface plays a key role in nutrient conservation and recycling. We hypothesize that: 1.) The root mat and humus layer on the forest floor act as an exchange column to prevent leaching of nutrients until the nutrients can be taken up by the roots. (2.) Mycorrhizal fungi play a role in the direct transfer of nutrients from decomposing litter to roots. Other nutrient conserving mechanisms that we are examining are: (3.) Algae and lichens living on the surfaces of leaves and bark play an important role in nitrogen fixation of the forest. (4.) There are virtually no nitrifying bacteria in the forest. Maintenance of nitrogen in the ammonium form is a nitrogen conserving mechanism (Rice & Pancholy, 1972). (5.) There is either a sulfur fixing capability in the forest such as sulfur fixing bacteria, or the forest is being depleted of sulfur, since loss of sulfur through stream flow far exceeds input through rainfall. (6.) Sclerophylly and evergreenness in the tropical rain forest are nutrient conserving mechanisms. 740 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 (7.) Many nutrients move from leaves back into the stems before the leaves fall. (8.) Trees are adapted to the oligotrophic environment in that roots are physiologically very efficient in extracting nutrients, utilize a low oxygen en- vironment, and, at least in the podsol site, are resistant to flooding. (9.) Insect predation of leaves is low in the podsol site, and only slightly higher in the laterite site. These low predation rates may be due to plant com- pounds such as alkaloids and polyphenols. These compounds may act as nutrient conserving devices in that it is more economical for the plant to manufacture secondary compounds than it is to manufacture a new leaf in the nutrient poor environment. (10.) Termites play an important role in redistribution of nutrients in the (11.) In the tierra firma site, the rough root mat on the soil surface causes the newly fallen leaves to lie at various vertical angles, with the result that the leaves resemble somewhat the shingles on a pitched roof. Rainfall and through- fall quickly pass over these “shingles,” minimizing the opportunity for leaching by water, and allowing more time for recycling by mechanisms such as mycor- rhiza. Other hypotheses are emerging relevant to the treated experimental areas: (12.) Despite the fact that the roots of secondary successional species are primarily in the upper layer of mineral soil and not on the soil surface, they have an extremely high capacity for nutrient uptake. When the forest is cut, but allowed to immediately begin the successional process, the successional species can recover a large proportion of the nutrients released by the decaying organic matter. However, if the ecosystem is cropped, most of the nutrients will be lost, either through leaching or through harvesting. 13 ife spans of slash and burn farms are determined primarily by the decay rate of organic matter and root biomass in the soil which supplies nutrients to the crops. In addition to development of these ideas, comparison of the data from the two forest types with different soil conditions has led to hypotheses regarding nutrient cycling in the podsol sites versus the lateritic sites, as well as hypotheses regarding cycling in these ecosystems compared to other forest ecosystems: (14.) In the laterite sites, standing crop, productivity, and rates of nutrient cycles are slightly higher than in the podsol, seasonally flooded site, possibly due to lesser extremes of water conditions and anaerobiosis. (15.) Highly sclerophyllous vegetation with highly inclined leaves located in patches on the podzolic soils, with a xerophytic aspect, reflect an extreme where there occurs drastic alternations between drought conditions and flooding with anaerobic conditions. (16.) Consumption of vegetation by insects in the podsol site is lower than in the laterite site. (17.) In both sites, rates of productivity and nutrient cycling are lower than on more fertile soils in both temperate and tropical regions. 1977] JORDAN & MEDINA—ECOSYSTEM RESEARCH 741 (18.) Although biomass in both sites is relatively low in comparison with other forests, the forests are climax in the sense that net ecosystem productivity is zero. (19.) The biomass of the forest is limited by the available pool of nutrients and the capability of nutrient-retaining mechanisms to prevent their loss. Other relationships that have emerged as a result of our studies of water balance and biomass, which were necessary steps im the quantification of the nutrient budget, are: (20.) Rate of transpiration in trees is independent of species and site, and depends only on sapwood area per unit of forest floor. (21.) Biomass of all tree species can be described by a single regression on (diameter)? (height) (density). ORGANIZATIONAL ASPECTS The San Carlos project is a cooperative study between institutions in Venezuela, the United States and Germany. The project is headquartered at Centro de Ecología, Instituto Venezolano de Investigaciones Científicas (LV.LC.), in Caracas, Venezuela. Other participating Venezuelan institutions are Universidad Central de Venezuela, and CODESUR, a branch of the Ministry of the Environment. The German participating institutions are the Max Planck Institute at Plón, and the World Institute of Forestry at Reinbeck (Hamburg). Participation of United States institutions is being coordinated through the Institute of Ecology, University of Georgia. Funds for the project are coming from the Organization of American States (OAS), UNESCO, CONICIT (Venezuelan Science Foundation), United States National Science Foundation, Deutsche Forschungsgemeneinschaft, and in- directly through IVIC and CODESUR. The project has been designated a MAB I pilot project by UNESCO be- cause of the progress that has been made in relation to other MAB ecosystem studies in the tropics. It is also part of the Humid Tropics Forest Project of the OAS, which includes projects in Brazil, Trinidad, and Colombia. The project was started in 1974, at just about the time the International Biological Program (IBP) Biome studies were drawing to a close. In designing the project, we strove to take advantage of the lessons learned during the operation of the IBP studies. The strengths and weaknesses of these programs have been discussed by Mitchell et al. (1976). In order to build upon the wisdom gained from the IBP studies, we did the following: (1.) First of all, we returned to the old-fashioned method of designing the project, to test hypotheses, rather than build the project around a technique, such as was done with systems analysis in the IBP studies. (2.) Secondly, we confined our ecosystem model to a single process model, rather than model many processes and populations and attempt to integrate them into a single model, as had been done with little sucess in the IBP studies. (3.) Thirdly, we kept the project small, in comparison with the United 742 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 | States IBP studies. In many cases, the large scale of these studies had caused them to be unwieldy from the point of view of management. Another organizational factor which weighs heavily in biological research in the tropics, and especially ecosystem research because of its magnitude, is the problem of scientific imperialism. For many decades, if not for centuries, North American, European, and Japanese scientists have visited Latin American, African, and Southeast Asian countries, collected data, specimens, and samples, brought them back to their home countries, and bestowed little or no scientific benefits upon the host countries. Over the years, this has resulted in a resent- ment in the tropical host countries because knowledge derived from such studies or whole efforts did not contribute to the improvement of personnel and infra- structure in the respective countries, and did not aid the development of similar projects run by their own people. Many times it has been due to lack of local scientific personnel, but often the projects did not have the policy of improvement of local capabilities ( Budowski This problem was discussed during the 1973 Costa Rican meeting of tropical ecologists, during which ecosystem research in the tropics was evaluated and recommendations for future research was discussed. The proceedings of this conference were later published in the book Fragile Ecosystems (Farnworth & Golley, As a result of the recommendations in this book, our ecosystem project was specifically designed to contribute to the scientific infrastructure of the host country. For example, instead of bringing samples back to the United States or Germany for analysis, we have set up our own analytical laboratory in Caracas, and trained a team of technicians to operate it. Part of the data processing is taking place in Caracas, but copies of all original data are kept in Caracas, so that it can be used by other investigators. We have limited the number of North American and European visitors, with the intention to increase as much as possible the number of Latin American participants. Further, we make an effort to have counterparts for visiting scientists, so that the visitors experience will not be lost to Venezuela. For ex- ample, we have initiated a soil microbiology program, in which the visiting United States microbiologist is traning a Venezuelan investigator to follow through and complete a study of nitrifying bacteria in the Amazon forest. COMPARISON WITH OTHER ECOSYSTEM STUDIES In general, most of the values of the ecosystem parameters which we have obtained so far are equal to or somewhat lower than values from other tropical ecosystem studies. Total living biomass on the tierra firma sites near San Carlos averaged 391 t/ha. Other biomass studies of tropical rain forests have produced values within the same range or somewhat higher. In Puerto Rico, Jordan (1971) esti- mated the biomass of one site of a montane forest to be 228 t/ha, while Ovington & Olson (1970) estimated three in the vicinity to be 324, 209, and 269 t/ha. Dry weight of above ground biomass in two Panama forests were 377 and 276 t/ha (Golley et al., 1975). In Ghana, Greenland & Kowal (1960) estimated 1977] JORDAN & MEDINA—ECOSYSTEM RESEARCH 743 biomass of a secondary forest to be 289 t/ha, while two evergreen tropical forests in the Ivory Coast, constituting part of the French project at the Banco and Yapo reserves, were estimated to have above ground biomasses of 465 and 425 t/ha (Huttell & Bernhard-Reversat, 1975). In the Pasoh forest of Malaya, above ground dry weight of biomass was 664 and 475 t/ha on two plots (Kato et al., 1974), considerably greater than our values for San Carlos. In evergreen seasonal forests of Cambodia, total biomass in two stands was 415 and 348 t/ha, (Hozumi et al., 1969), while in Thailand values ranged from 326 to 404 t/ha (Ogawa et al., 1965). Near Manaus, Brazil, Klinge & Rodriguez (1973) found about 900 t/ha fresh weight including roots. If we assume the moisture percentage is the same as in San Carlos, then total dry weight would be 585 t/ha. Rodin & Bazilevich (1967) in their survey of global biomass put an average value for tropical forests greater than 500 t/ha. The world biomass summary by Art & Marks (1971) gives similar high values. Leaf fall values which for the San Carlos forest are around 5 t/ha/yr are in the low part of the range of values for tropical forests. For example, in the Ivory Coast Forest Project, leaf fall rates were 8-10 t/ha/yr (Huttel, 1975) and in the Khao Chong forest, Thailand, rates were about 12 t/ha/yr (Kira et al., 1967). In the eastern Amazon Basin, near Belém, rates were 7.4-10.7 t/ha/yr, but in the central basin near Manaus, the rate was 6.7 t/ha/yr (Klinge, 1974), only slightly greater than the value for San Carlos. Jordan & Murphy (1977) have presented litter fall values from 27 tropical forests. Most values are greater than 7 t/ha/yr, and there are quite a few values greater than 10 t/ha/yr. Rates of soil respiration at San Carlos are about 400-500 mg C/m*/hr, within the range encountered in Thailand. Concentrations of nutrients in water fluxes such as throughfall, stem flow, and soil water are generally less than were found in tropical forests in Puerto Rico (Jordan, 1968), Panama (Golley et al., 1975), and Ghana (Nye, 1961). The most striking difference between our study, those of Klinge (1973), those of Went & Stark (1968)—all in the Amazon Basin—and those studies in other regions of the tropics is the apparent importance of the root mat and humus layer in the Amazon forests. As we mentioned previously, the root mat appears to play a key role in the recycling of nutrients. Yet in other studies outside the Amazon region, the presence of a surface mat, if present, is not noted or empha- sized. The evidence that we are obtaining, then, is verifying the idea that the Amazon forest is severely nutrient limited, and that low biomass, low litter fall, and low nutrient concentrations, all are adaptations to the oligotrophic condition. In addition, mechanisms such as the above-ground root mat and direct recycling by mycorrhiza are adaptations to help the ecosystem survive in the nutrient-poor conditions. The implication is that destruction of the Amazon forest on a large scale will cause an irretrievable loss of nutrients and conse- quently of the ecosystem, because large scale clearing destroys the nutrient con- serving mechanisms. 744 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 FUTURE TropicAL EcosYSTEM Work What about future ecosystem research in the tropics? It is typical for a presentation to conclude with a plea that it is especially important for the particular research discussed to receive more recognition and greater support. We will not break with this tradition. In general, ecosystem research in the tropics is too fragmented, with the result that the studies do not have the political effectiveness that they should, in the sense that the results of ecosystem research should influence political planning for a region. When a group of scientists work together on a single ecosystem problem, such as they did on the Hubbard Brook study (Bormann et al., 1968), the final impact is much greater, even if the findings are very con- troversial as they were in the Hubbard Brook study (Aubertin & Patric, 1974). For this reason then, we make the plea for less fragmented research and more integrated efforts, with the ecosystem approach being a natural integrating evice. LITERATURE CITED Art, H. & P. Marks. 1971. : summary e of biomass and net annual production in forest ecosystems of the world. Pp. 3-32, in Forest Biomass Studies. Proceedings of the XVth IUFRO Cae. College of Life nde and Agriculture, Orono, Maine a G. M. & J. H. Patric. 1974. Water yid after clear-cutting a small watershed West Virginia. J. Environm. Qual. 3: 243-249, 3 F. G. E. Likens, D. W. FisugR & D. PrEncE. 1968. Nutrient loss ac- celerated by dlear- cutting of a forest ecosystem. Science 159: 882-884. BupowsKi, G. Scientific Imperialism. Mimeograph. International Union for Conservation of ature and Natural Resources, Morges, Switzerland. Farnworth, E. G. & B. GorrEev. 1973. Fragile ern Evaluation of Research and Applications in the Neotropics. Springer Verlag, New York. GoLrEv, F. B., J. T. McGinnis, R. CLEMENTS, G. Currp pi M.J. Dvuever. 1975. Mineral Cycling ina pia Moist Forest Ecosystem. Univ. of Georgia Press, Athens, Georgia. = — GREENLAND, D. & J. Kowar. 1960. Nutrient content of the moist tropical forest of hana. Pl. & Soil 12: 154-1 0 Hozumi, K., K. Yopa, S. KORA & T. Km 1969. Production ecology of tropical rain forests in southwestern . d I. Plant biomass. Nature and Life in Southeast Asia 1—34. ron bet mie 1975. Recherches sur l'ecosysteme de la foret Scu ur de Basse Cote-d'Ivoire. V. Biomasse vegetale et productivite primaire cle de la matiere organique. Terre & Vie 29: -228. JonpaN, C. F. 1968. Concentration of elements in forest water. Pp. 47-50, in Puerto Rico uclear Center, The Rain Forest Project Annual Report PRNC-119. 1971. m of a tropical forest and its relation to a world pattern of energy EL us Ecol. 127—142. s ecology. Bull. Ecol. Soc. Amer. 56(2 MURPHY. 1977. A latitudinal gradient of wood per litter production, and its SER regarding competition and species diversity in trees. Amer. Midl. Naturalist in press . Unt. In preparation. Biomass of a “tierra firma" forest in the Rio Negro region of the 1 Basin. Manuscript. Karo, R., Y. Tapaxr & H. Ocawa. 1974. Plant biomass and growth increment studies in Pasoh forest. IBP synthesis meeting, Kuala Lumpur, 12-18 Aug. 1974. Mimeographed repor Kira, T., H. Ocawa, K. Yopa & K. Ocrvo. 1967. Comparative ecological studies on three main types of forest vegetation in Thailar id IV. Dry matter production with special reference to the Khao Chong rain forest. Nature and Life in S. E. Asia 5: 149-174. 1977] JORDAN & MEDINA—ECOSYSTEM RESEARCH 745 Kuınce, II. 1973. Root mass estimation in lowland tropical rain forests of Central Ama- zonian, Brazil. I. Fine root masses of a pale yellow latosol and a giant humus podzol. 38. 974. Litter production on tropical an IBP synthesis meeting, Kuala Lampur, August 1974. Mimeographed report 1976. Phytomass of dominant tree species in an Amazon caatinga. VII Symposium on , Biogeographical and Landscape-Ecological 1 in South America, May 1976 Plön, Germ & W rp iS 1973. Biomass estimation in a Central Amazonian rain forest. Acta Ci. Venez. 24: 225-237. MITCHELL, R., R. A. Mayer & J. DowWNJHOWER. 1976. An evaluation of three biome pro- grams. Science 192: 865. Nye, P. 1961. Organic matter and nutrient cycles under moist tropical forest. Pl. & Soil : 33 6. Ocawa, H., K. Yopa, K. Ocio & T. Kma. 1965. Comparative ecological studies on three main types of forest 5 in Thailand II. Plant Biomass. Nature and Life in Southeast 578 4; 49-80. Ovincton, J. D. & J. S. Orson. 1970. Biomass and chemical content of El Verde lower montane rain forest plants. e H-53-H-77, in H. T. Odum (editor), A Tropical Rain Forest. Div. Tech. Inform. U. S. Atomic Energy Commi ission PLATrr, R. 1974. Who paid for ecology. Bull. Ecol. Soc. Aot. 55(4 Rice, E. & S. PaNcHoLy. 1972. Inhibition of nitrification by climax B Amer. J. Bot. 59: 1033-1040. RobiN, L. E. & N. BAziLevicH. 1967. Production and Mineral Cycling in Terrestrial Vegeta- tion. 9 from Russian. Oliver and Boyd, London. 208 pp. Went, F. & N. Stark. 1968. Mycorrhiza. BioScience 18: 1035-1040. PERSPECTIVES IN TROPICAL BOTANY: CONCLUDING REMARKS PETER H. Raven! It would be unrewarding to attempt to summarize the rich sampling of per- spectives in tropical botany that has been provided by the five papers in this symposium. Each should be read for what it has to offer in insight and in vistas for future research. In these concluding remarks, I wish instead to offer some thoughts on the general state of tropical research, and to attempt to put them into a global perspective. udged from statistics about relatively well known groups of animals and plants, there are likely to be about twice as many species of any group in the tropics as in temperate regions. About 1.5 million kinds of organisms have been given names during the first 225 years of our effort to do this, and there are probably at least twice as many that remain to be named. Of these, perhaps two-thirds of the estimated 1.5 million organisms of temperate regions have been named, but no more than one in five of those in the tropics. In 1975, the following estimates of cumulative destruction of tropical forest were made by the Food and Agriculture Organization (FAO) of UNESCO (Sommer, 1976) : 9 Original area of tropical rain forest (world): 16 million km? Area in 1975: 9.35 million km? (reduction of 41.5% ) Destruction to 1975: Latin America 36.6% Southeast Asia 38.1% Africa 51.9% India, Sri Lanka, Burma 63.3% A doubling in population size is projected for all tropical countries by the end of the century, Although inroads are being made into the difficult problems of expanding food production in the tropics, the Environmental Fund predicts that annual population growth will outstrip growth in food production in each of the major tropical areas of the world through 1985, which is as far as they have attempted to project. Statistics such as these lead to forecasts such as that of the destruction of all moist tropical forest in the Philippines and Malesia within 5 to 10 years, and throughout Indonesia within 15 to 20 years. Worldwide, it appears likely that virtually all tropical forest will be destroyed or at least irreparably damaged by the close of the century, although it will probably still persist in a few local areas, perhaps including portions of the Amazon Basin, by that time. Projected rates of growth in population, food, and energy lend little hope for a reversal of these trends. For example, the FAO estimates that by 1985 some ‘Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110. ANN. Missouni Bor. Garp. 64: 746-748. 1977. 1977] RAVEN—CONCLUSIONS 747 26 countries, with an aggregate population of 365 million people, will be unable to produce enough food to prevent gradual starvation. In addition, if most tropical forest will have been removed by the end of the century, what of the quarter to half million people who make their living at present by slash-and-burn agriculture im this forest? Tens of millions of people in the world starve to death each year at the present time, and with these trends being what they are, it appears likely that hundreds of millions will starve in tropical countries during the coming two decades. The countries involved, which will mostly become preoccupied with life-and-death questions of food and energy, will be unlikely to be able to divert many of their precious financial reserves to the study of the forests and other natural ecosystems, even though the sustained productivity of their lands ulti- mately depends upon such knowledge (cf. Janzen, 1973). It therefore will ap- parently be up to developed countries such as the United States to devote the necessary capital to gaining whatever ecological knowledge it might be possible to accumulate during these critical years. Only by doing so will it be possible to gain a measure of world stability for our successors in the twenty-first century. In the light of these facts, it would appear that there is little or no chance for the long-term preservation of a significant sample of tropical diversity. Many of the two million or more species of animals and plants that occur in the tropics and which have never been catalogued or given a scientific name will become extinct before they are collected once; however, since our generation alone will be able to deal with them, we should make every effort to preserve samples for those who will follow us to study and to use to answer a host of questions about the diversity of life on earth that have not even been formulated yet. In terms of formal inventories, the rates of completion of tropical floras are such that, when compared with the rate of destruction of the forest themselves, they appear pathetically low. We are simply not devoting sufficient resources to these projects to allow for their completion in a timely fashion. It would be highly desirable to step up our input into them, especially by involving more of the citizens and institutions of the tropical countries themselves in the project and by securing adequate funding from whatever sources appear possible. No matter how energetic these efforts may be, however, it is clear that the inventories will not be completed in time to provide a basis for further biological generalities or extrapolations of higher order: questions about biological interactions must be asked now, if we are to learn anything about the functioning of ecosystems upon which a growing proportion of the human race depends for survival. Tropical biology of all kinds must receive a high priority in both research and training programs, at every level, and within every competent body. We can learn about the nameless and unknown plants and animals of the tropics only for a few more decades; our survival depends directly upon an understand- ing of the way in which they maintain a stable productivity in the hottest and rainiest areas of the world, often on extremely infertile soils. The attainment of such knowledge by our own generation is of crucial importance, since our children and grandchildren will not be able to help. 748 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 64 LITERATURE CITED JANzEN, D. II. 1973. Tropical agroecosystems. Science 182: 1212-1219. SOMMER, A. 1976. Attempts at an assessment of the world's tropical forests. Unasylva 28: 5-27. The previous issue of the ANNALS OF THE MISSOURI BOTANICAL GARDEN, Vol. 64, No. 3, pp. 381-655, was published 26 May 1978. > WRITE FOR -SQOMPLETE — .. 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