A Monograph of the Genus Myrceugenia (Myrtaceae) by Anne Barber

Organization for Flora Neotropica

A Monograph of the Genus Myrceugenia (Myrtaceae) Author(s): Leslie R. Landrum Source: Flora Neotropica, Vol. 29, A Monograph of the Genus Myrceugenia (Myrtaceae) (Dec. 11, 1981), pp. 1-135 Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica Stable URL: http://www.jstor.org/stable/4393751 Accessed: 03/05/2010 12:43 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=nybg. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.

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A MONOGRAPH OF THE GENUS MYRCEUGENIA (MYRTACEAE)

LESLIE R. LANDRUM

CANCER Of

TROPIC

FLORA NEOTROPICA, TiROiC

CAPRICORN

/ Xy

FLORA NEOTROPICA MONOGRAPH Number 29

The New York Botanical Garden Bronx, New York 10458 Issued II December 1981

This materialis based upon research supportedby the National Science Foundatonunder DEB 76-21543and is published with support of National Science FoundationGrant No. DEB 80-05535. The Foundation provides awards for research and education in the sciences. The awardee is wholly responsible for the conduct of such research and preparationof the results for the publication. The Foundation, therefore, does not assume responsibility for such findings or their interpretation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author and do not necessarily reflect the views of the National Science Foundation. Library of Congress Cataloging in Publication Data LANDRUM,LESLIER., 1946-

A monographof the genus Myrceugenia (Myrtaceae) (Flora neotropica ; monographno. 29) Based on the author's thesis (Ph. D.)University of Michigan, 1980. Bibliography:p. Includes index. 1. Myrceugenia. 2. Botany-South AmericaClassification. I. Title. II. Series. QK495.M9L36 1981

ISBN 0-89327-236-1

583'.42

81-11282

AACR2

Published by The New York Botanical Garden Bronx, New York 10458

International Standard Serial Number 0071-5794

All material subject to this copyright may be photocopied for the non-commercial purpose of scientific or educational advancement.

A MONOGRAPH OF THE GENUS MYRCEUGENIA (MYRTACEAE) LESLIE R. LANDRUM'

TABLE OF CONTENTS I. Introduction ....................... II. Taxonomic History ................ III. Position of Myrceugenia in the Myrtaceae ...................... IV. Phylogeny and Geography .......... V. Systematic Criteria ................ 1. Habit ......................... 2. Hairs ......................... 3. Leaves ........................ 4. Inflorescence .................. 5. Flowers ....................... ....... 6. Fruit ................. 7. Phenology ..................... 8. Wood Anatomy ................ 9. Chromosome Numbers ......... VI. Systematic Treatment . ...... ...... Key to the Species of Myrceugenia .. 1. M.fernandeziana .............. 2. M. rufescens .................. 3. M. campestris ................. 4. M. schultzei ................... 5. M. exsucca .................... 6. M. lanceolata ................. 7. M . rufa ....................... 8. M. ovata ...................... 9. M. pinifolia ................... 10. M. obtusa ..................... 11. M. parvifolia .................. 12. M. leptospermoides ............ 13. M. hatschbachii ............... 14. M. smithii ..................... 15. M. planipes ...................

I 2 4 8 10 10 10 10 13 14 16 16 17 18 19 22 30 32 33 34 35 37 40 42 47 49 50 51 52 54 56

VII. VIII. IX. X. XI.

16. M. kleinii ..................... 17. M. reitzii ...................... 18. M. correifolia .................. 19. M. chrvsocarpa ................ 20. M. glaucescens ................ 21. M. scutellata .................. 22. M. cucullata ................... 23. M. seriatoramosa .............. 24. M. colchaguensis .............. 25. M . franciscensis ............... 26. M . alpigena ................... 27. M . bracteosa .................. 28. M. euosma .................... 29. M. acutiflora .................. 30. M. venosa ..................... 31. M. myrcioides ................ 32. M. pilotantha .................. 33. M. miersiana .................. 34. M. mrtoides .................. 35. M. hoehnei .................... 36. M. brevipedicellata ............. 37. M. oxysepala .................. 38. M. leptocalyx .................. Excluded species .................. ................ Hybrids . ............ Acknowledgments . Literature Cited . ............. Numerical List of Taxa .......... List of Exsiccatae . ............ Index of Scientific Names .........

58 58 61 62 63 71 72 73 74 75 76 81 83 86 88 91 95 99 100 103 104 105 106 107 108 124 124 126 127 132

I. INTRODUCTION Myrceugenia is one of about 40 genera of the American Myrtaceae belonging to the tribe Myrteae, the fleshy fruited Myrtaceae (McVaugh, 1968). The species of Myrceugenia are trees and shrubs that grow in moderately cool, wet climates in temperate and subtropical South America. There are two species on the Juan Fernandez Islands, which lie about 700 km west of the coast of central Chile; 12 in central and southern Chile and adjacent Argentina; and 25 in eastern South America, the majority of these last being found along the eastern edge of the planalto and along the coast of Brazil from near Rio de Janeiro to Porto Alegre. A distribution map of the genus is shown in Fig. 1. The genus has been the subject of fairly recent taxonomic treatments. Kausel (1942, 1944, 1948, 1949) and Navas (1970) have considered the western species and Legrand (1957, 1961) and Legrand and Klein (1970) have done extensive B. A. Krukoff, Research Associate, The New York Botanical Garden, Bronx, New York 10458.

Flora Neotropica

studies of the eastern species. Yet, until now, no one has done a systematic treatmentof all the species. This study was undertakenwith the expectation that by considering both eastern and western species, a better taxonomy could be constructed and that an understandingof the taxonomy and distributionof the genus might lead to interestingphytogeographicinferences. One of the intriguingproblems associated with Myrceugenia is its anomalous position among the genera of the tribe Myrteae. It has an embryo like other members of the subtribe Myrciinae, viz. Myrcia, Calyptranthes, Gomidesia and

Marlierea;but in floralcharactersand those of the inflorescenceit is more similar to genera of other subtribes. Berg gave Myrceugenia its name because it is superficiallysimilarto Eugenia but has the embryo of Myrcia. My studies of Myrceugenia were done as a graduatestudent at the University of Michigan under the direction of Dr. Rogers McVaugh. The results first appeared as a doctoral dissertation (Landrum, 1980). For publication they have been divided into two parts: 1) the taxonomic treatmentof the genus which is the subject of this paper;and 2) the phylogeny and geographyof Myrceugeniawhich will appear as a separate paper (Landrum, 1981), but which will be summarized briefly below. II. TAXONOMICHISTORY During the early taxonomic history of the Myrteae, before the work of A. P. de Candolle and Berg, the generic limits as we recognize them today were not well understood. Genera were based on what now seem ratherarbitrarilychosen features such as the numberof calyx-lobes, locules or seeds. These characteristics where used today have been correlatedwith other characteristics. With the contributions of de Candolle (1828) and Berg (1859), the generic limits began to become clear. These workers recognized several genera that are now known to have a strong taxonomic basis. But because the Myrteae are so uniform,it was often difficultfor early students of the family to assign individual species, even to genera they themselves had described. Thus Berg, the authorof Myrceugenia, described from flowering specimens several new species that he assigned to Eugenia, althoughstudy of fruitingmaterialhas subsequentlyshown their correct position in Myrceugenia. It has been the task of more recent botanists to place many known species into the right genera. For Myrceugenia this work was mainly done by Legrand and Kausel between 1940 and 1953. The taxonomic history of Myrceugenia has been complicatedby the fact that Burrettook up the name Luma A. Gray (1854) in preference to Myrceugenia Berg (1856). In a series of two articles (1941a, 1941b)he apparentlytransferredevery species of Myrceugeniaknown to him to Luma and described several new species under that name. Luma (=Myrceugenella), as understoodby Kausel (1942), McVaugh(1968)and me, is a smallgenus, distinct from Myrceugenia.

Fortunatelyfor this taxonomic study, it is probablethat most species of Myrceugenia have been discovered and are recognized as belonging to the genus. The Myrtaceaeare fairly well understoodin all the regions in which Myrceugenia is known to grow and also in tropical America north of the known range of the genus. Because most species of Myrceugenia grow in areas that were accessible to the botanical explorers of the nineteenthcentury, the majoritywere describedat least once by 1900, and were usually assigned to Eugenia or Myrtus. Then taxonomic activity in the genus was low until about 1940when Legrandand Kausel

Myrceugenia

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FIG. 1. Distribution of the genus Myrceugenia. Each dot represents at least one collection. of Berg and Burret were housed there and it has often been impossible as yet

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began to transferspecies from other genera to Myrceugenia and to describe new taxa. Recently extensive botanical exploration has been done in the Brazilian states of Paranaby Dr. G. Hatschbach(of Curitiba)and Santa Catarinaby Doctors R. Reitz and R. Klein (of Itajaf) and L. B. Smith (of Washington, D.C.). Their abundantcollections have included several new taxa, most of which Legrand has described. The destruction of the Berlin herbariumduringWorld War II had an unfortunate effect on the taxonomy of Myrtaceaein general. Numerous type specimens of Berg and Burret were housed there and it has often been impossible as yet to locate duplicate specimens.

Flora Neotropica

4 h

III. POSITION OF MYRCEUGENIA IN THE MYRTACEAE The Myrtaceae have traditionally been divided into two groups based on fruit type. Niedenzu (1893) recognized these as subfamilies: the Leptospermoideae with dry, indehiscent or capsular fruits; and the Myrtoideae, with fleshy fruits. Kausel (1956) recognized the dry-fruited species as a separate family, the Leptospermaceae. The Leptospermoideae (or Leptospermaceae) are restricted to the Old World except for one monotypic genus, Tepualia which grows in the rain forests of southern South America. The Myrtoideae, which comprise one tribe, the Myrteae, are mainly American, but several genera and several hundred species are also found in the Old World. Myrceugenia is one of the genera of the Myrteae that is restricted to the New World. Basically two opinions have been expressed as to the position of Myrceugenia among the genera of the tribe Myrteae: that of Berg (1855) with which Kausel (1956) and Briggs and Johnson (1979) seem to concur; and that of McVaugh (1968). I will first discuss the opinions of Berg and McVaugh and then I will propose

my own, admittedlyspeculative, hypothesis. A more thorough,objective analysis

cannot be done until the genera of the Myrteae are better understood.

Myrceugenia Pimenta

Calyptranthes

PPsudocrryophyllus

Morlierea

Myrcia Gomidesia

/ 1/ /

Nothomyra

Hexachlamyr s

Myrceugenia

Eugenia

egrndia

Plinia

AeumyrtI

Siph ousutpn.

Myrciaria

I Calyrelpus

Amemyrtus

iatiOs

Myirtu

iv! CAnmprmnanesiy

>peproceiyx

/ -

Myrteolo

Psidium

Calycorectes

Uni

4 -l^ J -yrrhinium

t;itao

\ \

Tema

.P'./dnmomis

FIG. 3. McVaugh'sinformalgroups within the Myrteae. "Genera of the American Myrtaceae, tribe Myrteae,arrangedaccordingto their supposedevolutionaryaffinities.The circle includesthose generathat seem not to have been involved in any of the majorlines of descent . .."(1968, p. 371.).

Ever since the time of de Candolle (1828) it has been known that there are basically three types of embryos in the Myrteae. Berg used these as the definitive characters of three subtribes which in accordance with the modern rules of nomenclature would have somewhat different names than the ones he used. Thus Berg's Myrcioideae is now called the Myrciinae; his Eugenioideae, the Eugeniinae; and his Pimentoideae, the Myrtinae. The embryo of the Myrciinae (the subtribe to which Myrceugenia has traditionally been assigned) has thin membranous cotyledons folded into a bundle and a long horseshoe-shaped hypocotyl curled around the cotyledons (Fig. 2A, B). The embryo of the Eugeniinae is much like that of a bean, with a short hypocotyl and thick planoconvex cotyledons which may be either free or fused together (Fig. 2E, F). The embryo of the Myrtinae, often called pimentoid for historical reasons, has a relatively long hypocotyl and short cotyledons, and the whole is spiral, circular, hook-shaped or horseshoe-shaped (Fig. 2G). Berg also recognized two monogeneric subtribes based on other characters. These genera are probably most closely allied to genera of the Myrtinae. Berg more or less arbitrarily placed Myrceugenia in the subtribe Myrciinae along with Myrcia, Gomidesia, Calyptranthes and Marlierea because it has the same type of embryo as these other genera. In other characteristics it is quite different. McVaugh in his recent treatment of the American Myrtaceae rejects the concept of formal subtribes based primarily on the embryos. Instead he recognized

Flora Neotropica

six main lines of evolution (see Fig. 3) of which three would be included in the original subtribe Myrtinae(groups 4, 5 and 6); two in the Eugeniinae (groups 2 and 3); and one in the Myrciinae(group 1). In addition to the charactersof the embryohe used characteristicsof the seed coat, flower and inflorescenceto define these six informalgroups. He also identified eight genera which do not fit well into any of the six main groups. He considers these to be separate, independent lineages which have arisen from the same protomyrtaceousstock as the six main lines, but which have been adaptivelyless successful. He does not envision them as being ancestralto the six lines. Myrceugenia, and Nothomyrcia which I consider to be a synonym of Myrceugenia, are two of these eight genera. As was mentionedabove, Myrceugeniashares the myrcioidembryo with Myrcia, Calyptranthes, Marlierea and Gomidesia, members of McVaugh's group 1

and Berg's Myrciinae, but otherwise is different from them. Myrceugenia has 4-merousflowers instead of primarily5-merousflowers; it has simple, uniflorous, dichasial or bracteate shoot inflorescences instead of panicles; it has numerous ovules in each of 2-4 locules instead of 2 ovules in each of 2 (rarely 3) locules. In all these characteristics Myrceugenia resembles members of the subtribes Eugeniinaeand Myrtinae. If it were known that the three types of embryos had each arisen only once, then the subtribes as set down by Berg would be acceptable as monophyletic units. At least tentatively, I would like to assume that this is known and offer the following two reasons for believing it may be true. First, the differencesbetween the types of embryos are not simple but ratherinvolve significantchanges in the fleshiness of the cotyledons as well as in the length and conformationof the radicle. Secondly, the types of embryos are not plastic characters. I know of no genus of the Myrteae in which the embryo is variable, and only the genus Luma possesses embryos intermediatebetween two of the three types (Fig. 2C, D). If for argument'ssake we assume the three types of embryos have arisen only once and that the different subtribes defined by them are monophyletic, an interesting pattern emerges: within each subtribethere is at least one genus which has the following characteristics. 1. 2. 3. 4. 5. 6. 7.

No fusion of calyx-lobes beyond the top of the inferiorovary. No fusion of the cotyledons. An indefinitenumberof ovules in each locule of the ovary, usually 5-20. 2-4 locules per ovary, usually 2. Most or all species with 4-merousflowers. Seeds with membranoustesta. A stronglydeveloped dichasiumin at least some species.

Luma, the genus which has an embryo more or less intermediatebetween the eugenioid and myrcioidtypes, also has all these characteristics. It is probable that the ancestral group which would link the three subtribes togetherhad this set of characteristics.I provisionallyaccept these characteristics as being primitiveand the genera which have them as being similarto the ancestors of each subtribe. I would rathernot speculate as to which subtribeis the most primitive. But it is probably not the Myrciinae because they are restricted to the New World whereas the Myrtinae and Eugeniinae are widespread in both Old and New Worlds. Within the Myrtinae, the genus that is postulated as being most primitive is Blepharocalyxwhich is found in the Andes of Ecuadorand in temperateeastern South America. A Chilean genus Temu is perhaps not separable from Blepharocalyx (McVaugh, 1968).

Myrceugenia

f-MlSTfTI

9P.T

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*InEUb-V llwll

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FIG. 4. Postulatd position of t

FIG. 4.

:'in

the

e Postulated position of Myrceugenia in th tribe

Myrteae.

In the Eugeniinae, the genus postulated to be most primitive is Myrcianthes which grows in temperate eastern South America, in the Andes of Peru northward to Central America and in the West Indies. The Chilean genus Reichea is probably closely related (McVaugh, 1968). In the Myrciinae the postulated primitive genus is Myrceugenia found in temperate eastern and western South America and the Juan Fernandez Islands. Luma is known from temperate western South America and from a few localities in Peru and Bolivia. The fact that all these genera have widely disjunct distributions supports the idea that they are old. All have mainly temperate South American or Andean distributions. The cool climates of the Andes are probably not old because the mountains began their major uplift in the late Miocene (Rutland et al., 1965). Temperate South America may have served as a source of colonizers for montane Andean regions. Thus I believe that the American Myrtaceae probably originated in southern South America, a hypothesis which has also been suggested by McVaugh (1968). As will be discussed in the section on wood anatomy, it has been found that Myrceugenia and Luma have some vessels with scalariform perforation plates and that Blepharocalyx, Luma and Myrceugenia have tracheids. These structures have only rarely been found in the rest of the Myrtaceae and are generally accepted as being primitive. Their presence here supports the idea that these genera are evolutionarily not distant from one another and that perhaps they are primitive. Thus I propose that Myrceugenia may be similar to the ancestor of the Myrciinae and that it may also be similar to the ancestral group which would link the three subtribes together (Fig. 4).

Flora Neotropica IV. PHYLOGENY AND GEOGRAPHY

As mentionedearlierthe subjects of phylogeny and geographyhave been treated separately (Landrum, 1981) and the reader should consult that paper for a more thorough account. I will only attempt to summarizethe reasoning and conclusions here. In order to obtain objective estimates of evolutionary history two numerical methods were used: one based on the criterion of parsimony (Kluge & Farris, 1969)and another based on the criterionof compatiblecharacters(Estabrooket al., 1977; Estabrook & Meacham, 1979). A third analysis was done using the criterionof charactercompatibilityand charactercorrelationtogether. The results of these analyses were hypothetical undirectedphylogenetic trees. These trees served as an objective way of estimatingthe numberof transcontinentallinks in Myrceugenia. Because none of these numericalmethods took into account intraspecificvariationand possible or known cases of hybridization,a finalestimate was made which did include them as sources of data. This hypothetical phylogenetic tree, which represents the author's best estimate of evolutionaryhistory has been presented in Landrum(1981) and is repeated here (Fig. 5). The widely disjunct populations of Myrceugenia (Fig. 1) might be explained by two alternative hypotheses. The first, that continuous migration was once possible between the present areas of distribution,implies that an area that is not now supportingthe plants once was adequatefor their growth. An environmental change is therefore indicated. The second hypothesis is that long-distance dispersal is or was possible between the areas of distributionacross the barrier separatingthem and that the plants of one area could have become established after arrivingat the second area. In the case of the species of Myrceugenia on Juan Fernandezthe evidence is strongly in supportof the second hypothesis, essentially because continuous migrationhas most likely never been possible between the mainlandand the islands. According to Baker (1967) the islands are remnantsof volcanic cones and have probablyemerged from the ocean. On the other hand, the first hypothesis is consideredto more accuratelyexplain the continental disjunction of Myrceugenia because of (1) the apparentlypoor adaptationof the fruits and seeds to being dispersed over long distances; (2) the adaptationof the species to specific stable habitats which would make establishment in a new region difficult;(3) the presumedevolutionaryrelationshipsof the species in the genus; and (4) the distributionof other temperateSouth American genera. A continuous population of Myrceugenia is hypothesized to have existed across South America during the Tertiary, perhaps as early as the Eocene and as late as the beginningof the Miocene. Duringthis period oceanic transgressions covered southern Argentina and Chile (Fergulio, 1950; Harrington, 1962), the temperatureswere warmer than at present, and rainfall was heavier in central Argentinathan at present (Volkheimer, 1971). Thus a climate similar to that in which Myrceugenia now grows probably existed in central Argentina, i.e. one with an averageannualrainfallbetween 1-3 m and an averageannualtemperature between 10 and 20 degrees centigrade. Myrceugenia probably became divided into eastern and western populationsduringthe Miocene due to climatic changes caused by the rise of the Andes and the disappearanceof the oceanic transgressions.

Ukr

~~~S~~~c,IL~~~~~

FIG. 5. Postulated phylogeny of Myrceugenia. This estimate of evolutionary history is based on numerical intraspecific variation. Each taxon is represented by a three letter symbol, the first two letters are an abbre indicates if the species is eastern or western (ACe-M. myrcioides var. acrophylla; AFe-M. acutiflora; ALealpigena var. rufa; ATe-M. ovata var. acutata; BEe-M. brevipedicellata; BTe-M. bracteosa; CAe-M. cam M. colchaguensis; CRw-M. correifolia; CUe-M. cucullata; EUe-M. euosma; EXw-M. exsucca; FEw-M. var. glaucescens; GRe-M. ovata var. gracilis; HAe-M. hatschbachii; HOe-M. hoehnei; KLe-M. kleinii; L var. longifolia; LSw-M. leptospermoides; LTe-M. glaucescens var. latior; LXe-M. leptocalyx; MAe-M. cioides var. myrcioides; MIe-M. miersiana; MYe-M. myrtoides; NAw-M. ovata var. nannophylla; OBw-M OXe-M. oxysepala; PAw-M. parvifolia; PIw-M. pinifolia; PLe-M. pilotantha var. pilotantha; PPw-M. rufescens; RUw-M. rufa; SCw-M. schultzei; SLe-M. scutellata; SMe-M. smithii; SRe-M. seriatoramosa

Flora Neotropica V. SYSTEMATICCRITERIA Habit

The species of Myrceugeniaare all woody, varyingfrom small much branched shrubs less than 1 m high to trees over 10 m high with one central trunk. Most commonly the species are shrubsabout 2-5 m high. In general, species that seem to be closely related on other morphologicalgrounds have similar habits, but there are exceptions to this rule. Little use was made of habit in these studies, but it will sometimes prove to be a good field character(Fig. 6). I have never tried to ascertain the age of an individualof Myrceugenia, but I suspect that some species often live for several decades because they are members of stable, persistent communities. Many species are able to regenerateby means of stump sprouts, so one genetic entity might persist in this way indefinitely. Hairs The hairs of Myrceugenia are of great taxonomic importance in their form, density, and location. Nearly all species have dibrachiatehairs which perhaps cannot be said of any other genus of equal size in the Myrteae, although most genera have some, and sometimes they are abundantand conspicuous. In M.

fernandeziana only simple hairs have been found, and M. miersiana and M.

rufescens generallyhave only simple hairs, but all the other species have at least some dibrachiatehairs. The dibrachiatehairs rangefrom symmetricalto strongly asymmetrical,with one arm many times longer than the other. The hairs may be of one type or of differenttypes intermingled,or the type may vary from one part of the plant to another. One common combinationis for the hairs on the twigs to be simple or asymmetricallydibrachiateand spreading,while the hairs on the leaves are symmetricallydibrachiate,or nearly so, and appressed (Fig. 7A, B). The location and density of the hairs is probablyone of the best indicatorsof systematic position. Some species are densely covered with hairs on the hypanthium, petals, calyx-lobes, bracteoles, peduncles, lower leaf surface, and twigs. At the other extreme, there are species that possess hairs only on the hypanthium or practically none at all. A general classification of the species has been constructed (Table I), using the location of dense hair cover as a basis. I consider "dense" to mean that about 40 percent or more of the underlying surface is covered by hairs and "sparse" to mean that no more than about 10 percent is covered. It should be rememberedthat in Table I the evaluations of each species are idealized and that individualspecimens may not conform exactly. Leaves

The leaves of Myrceugenia (and the Myrteae in general) are decussate (occasionally ternateor suboppositeon some twigs) and perennial,with entire margins. They vary greatly in size, shape and texture. They may be less than 1 cm long or over 10 cm long, narrowlylinear, elliptic, oblanceolate, obovate, lanceolate, ovate, or nearly orbicular.The texture varies from stiffly coriaceous to more or less membranous.The relative prominence of the veins is sometimes a useful characteras is the occasional presence of a mucro. The petiole is normallychanneled adaxiallyin species with largerleaves, but in one group the petiole is terete or nearly so. The base and apex may be acute, bluntly or sharply acuminate, obtuse, or rounded. The base can also be cuneate and is occasionally cordate. Narrow leaves are common among species which grow along streams so there

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FIG. 6. Selected habitats of Myrceugenia. A, rain forest at Paranapiacaba, ca. 30 km east of Sao P central Santa Catarina, Brazil; C, eastern extremity of the planalto at Fortaleza near Cambara, Rio Gr Mirasol ca. 36 km north of San Antonio.

Flora Neotropica

;~ tk,%,;~-~";~,:

.....~ .~

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: ...:fZ!:,j "'r~~YI~~L;"~.-. L.b~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ :

FIG 7. SEM photomicrographsof hairs, leaf surfaces, and wood. A, Myrceugeniaobtusa, hairs

on the hypanthium (Hutchinson 193); B, M. schultzei, hairs on peduncle (Skottsberg s.n.); C, M. lanceolata, lower leaf surface covered with flakes of wax (Claude Joseph 2913); D, M. euosma,

lower leaf surfacewith recessed stomata(Smith & Klein 10564);E, M. oxysepala, ring-likestructure surroundingthe stomata(Reitz 6492);F, M. lanceolata, simpleperforationplate and G, a scalariform

perforation plate in the same specimen (Rusbv 584); H, M. euosma, spiral thickenings in vessels (Legrand 754).

may be some adaptive value in having narrow leaves in that habitat. As would

be expected, strongly coriaceous leaves are found in species judged on other grounds to be xerophytic. In a survey of the lower leaf surfaces of Myrceugeniausing the scanningelectron microscope, I found a few interesting characteristics (Fig. 7C-E). Some species usually have a ring-like structure, apparentlya manifestationof the accessory cells, surroundingthe stomata while others do not have this structure. In M. M. alpigenta and M. M. euosma euosma the the stomata stomataare are generally recessed among alpigena and generallydeeply deeply recessed among In

Myrceugenia Table I

Distributionof Dense Hair Cover Outer petal surface

Outer calyxlobe surface

Outer bracteole surface

Peduncles

2 3 4 5 6 7 8

-+ -

10 11 12

Type

+ ?

+ +

+or -

+ or

Twigs

+ + + + + or + +

+ + + + + or +

+ or -

+ or + + or+

Lower leaf surface + +

+ -

+ = dense cover sparse to glabrous + = intermediate or variable

Species with Each Type of Distributionof Dense Hair Cover 1-M. acutiflora, M. miersiana, M. myrcioides, M. myrtoides, M. pilotantha, M. venosa 2-M. alpigena, M. bracteosa, M. colchaguensis, M. euosma, M. franciscensis 3-M. rufa 4-M. brevipedicellata, M. hoehnei 5-M. leptocalyx, M. oxvsepala 6-M. exsucca, M. lanceolata, M. reitzii, M. rufescens, M. seriatoramosa 7-M. correifolia, M. schultzei 8-M. cucullata 9-M. campestris, M. kleinii 10-M. fernandeziana, M. leptospermoides, M. obtusa, M. ovata, M. pinifolia, M. planipes, M. smithii Type 11-M. hatschbachii, M. parvifolia Type 12-M. chrysocarpa, M. glaucescens, M. scutellata

Type Type Type Type Type Type Type Type Type Type

the epidermalcells. A few species may have abundantflakes of wax covering the surface. All these characteristicswere sufficientlyvariablewithin species, so that a more extensive survey would have been necessary in orderto make them useful in this study. But I think that they may prove helpful in the future for solving certain difficulttaxonomic problems, such as suspected cases of hybridization. Inflorescence Briggs and Johnson (1979)have recently publishedan extensive analysis of the myrtaceous inflorescence with the objective being to elucidate evolutionary trends in the Myrtaceae. They introduce several new terms, some of which are replacements for older, traditionalones. Perhaps this is necessary in an evolutionary analysis but adoptionof their largelyunknownterminology,which seems at times excessively specific, would reduce the usefulness of this taxonomic study. Therefore I have chosen to utilize a descriptive language that has traditionally been used by students of the Myrtaceae, except that I have used the name "bracteate shoot" for what has generally been called a raceme.

Flora Neotropica

The simplest myrtaceousinflorescence, but not necessarily the most primitive, is a solitary flower subtended by a normalleaf. Myrceugenia ovata and several related species have this type of inflorescence. Three kinds of elaborationof this simple inflorescence are found in Myrceugenia (Fig. 8). The first type occurs when secondary flowers arise from the axils of the two bracteoles found at the base of every hypanthium,resulting in a three flowered dichasium. Flowers may also arise from the axils of the bracteoles of the secondary flowers to form a dichasiumof more than three flowers (Fig. 8C). The second type of elaborationoccurs when the leaves subtendinguniflorous peduncles are reduced to bracts (morphologicallysimilar to the bracteoles) on the branchlet. This branchletis here called a bracteate shoot and may terminate in a flower or in a vegetative bud. Briggs and Johnson (1979) have called the former a "botryoid" and the latter a "botryum." In reality a bracteate shoot is a compound inflorescence, i.e. a raceme of uniflorousdichasia (Fig. 8A). The third type of elaboration, common in Myrceugenia, is the groupingtogether of peduncles in a row in the axils of leaves or bracts and the grouping together of branches of the dichasiumin the axils of bracteoles. Often two, but as many as four or five peduncles may be verticallysuperimposedin a file between the shoot and the petiole or the bract (Fig. 8E). This peculiar form of massing togetherof flowers is found to a much lesser degree in other generaand combined with other floral characters can usually be used to identify Myrceugenia when fruits are lacking. When the leaves of a whole branchingsystem are reduced to bracts, the result is a panicle that may include bracteate shoots and dichasia as subunits. Such an inflorescence does not occur in Myrceugenia but is often found in Myrcia and other genera of the Myrciinae. In general, it can be said that in Myrceugenia, inflorescence size is correlated with leaf size. Thus dichasia, bracteate shoots and rows of three or four flowers are usually found in larger-leavedspecies and solitaryflowers or pairs of flowers are found in smaller-leavedspecies. It may be supposed that a certain amountof massing together of flowers is necessary to attractpollinators;thus larger-leaved species, in which the internodes are relatively long, must have several flowers together at each leaf axil in order to produce an adequate display whereas this is unnecessaryin smallleaved species because the internodesare generallyshort. Flowers The calyx-lobes are perhaps taxonomicallythe most useful floral structurein the Myrtaceaeand have proven to be so in Myrceugenia. They vary in size and shape (Fig. 8), being at one extreme broader than long, rounded, and strongly concave; and at the other extreme, longer than broad, acute, and scarcely concave at the base or not concave at all. In M. myrcioides acute calyx-lobes have become valvate but in the other species the calyx-lobes are imbricate to some degree. The bracteoles vary as much as or more than the calyx-lobes (Fig. 8). They are normally lanceolate to ovate but may also be spatulate, linear or orbicular. In Myrceugeniafernandeziana they are caducous at about anthesis but in other species they persist, usually until the fruit is mature. The texture varies from membranousto coriaceous. The hypanthiumvaries somewhat in form from more or less globose to narrowly obconical but shape seems to be variablewithin species and hardto quantify. The length of the hypanthiumcomparedto the length of the calyx-lobes and bracteoles is sometimes a useful character. Normally the hypanthiumdoes not

Myrceugenia

i"~ ;)

FIG. 8. Flowers and inflorescencesof Myrceugenia. M. rufescens: A, bracteateshoot, 1.5x; B, flower, 5x (Pickel 5374). M. exsucca: C, dichasial inflorescence, 1.5x; D, flower, 5x (Maldonado 21). M. alpigena var. rufa: E, 4 superimposed peduncles, 1.5x; F, flower, 5x (Smith & Reitz 14281). M. myrcioides var. acrophylla: G, closed flower bud with valvate calyx lobes, 5x (Hatschbach 10972); H, fruit, Ix (Klein 567). M. pilotantha var. major: I, flower, 5x (Reitz & Klein 8093). M. oxysepala: J, flower, 5x (Reitz & Klein 8673). M. planipes: K, flower, 5x (West 4861). Drawn by Mrs. Karen Douthit.

Flora Neotropica

surpass the top of the ovary but in Myrceugeniaplanipes it is usually prolonged about 1-1.5 mm beyond the ovary. The stamens vary in numberfrom about 40 to over 500. In order to count them it is best to use a bud which is just at the point of opening. In flowers which have lost the stamens it is often possible to count the scars on the staminal ring. Althoughit is a rathertedious task to evaluate this character,stamen numberhas been of value in hybrid studies. The numberof ovary-loculesvaries from2 to 4 and is usually correlatedwith the size of the flower. Withineach locule are from two to about 20 ovules, which are normallyarrangedin two longitudinalrows. The numberof ovules is never stable in a species, nor even in the locules of a single ovary. Thus Myrceugeniadiffers from Myrcia and other members of the Myrciinae, which normally have two ovules per locule. In Myrceugeniathe absolute lines between locules are apparently sometimes blurredduringdevelopment so that some will occasionally have about as many ovules as would one and a half locules normally;then the ovules are not arrangedin two neat rows. The total numberof ovules in an ovary can be over 30 but no more than a few seeds (1-5) normallydevelop. Perhaps some ancestor of Myrceugenia developed all or nearly all its ovules into seeds. The peduncles may be lacking or up to ca. 3.5 cm long. They are usually flattenedand can be as wide as 2 mm at about 1 mm below the bracteoles. In one form of Myrceugenia ovata var. gracilis the peduncles are only about 0.2 mm wide. Fruit The fruits of Myrceugenia are berries. Little is known about them because (1) they often ripen in the winter months from June to Septemberwhen few collections are made; (2) they are not preservedwell in pressed specimens; and (3) they are often attacked by insects and fungi. One fruit attackingfungus on Myrceugenia alpigena var. rufa has kindly been identifiedby Dr. C. T. Rogerson of the New York Botanical Gardenas a memberof the Hypocreales. A study of these pests might prove interesting because they seem to be capable of causing the fleshy tissues of the ovary to develop almost normally while the ovules within are damagedby insects or are stunted by fungi. Such fruits never ripen. The color of the fruit as far as known is orange, red, or purple, but little is known of the intraspecificvariationof this character. Future collectors and field workers are urged to take careful note of fruit color. Phenology There are species of Myrceugenia flowering in every month of the year, but most flower during the summer and autumn from December to May. These species generally maturetheir fruits 9 to 12 months later in the summeror fall of the following growing season. Thus, it is common to find fruits and flowers on the same plant. A relatively small numberof species flower duringthe late winter and spring from August to November. These species sometimes mature their fruits rapidly within a month or two of flowering(M. campestris and M. rufes-

cens) or after a period of three to five months (M. rufa, M. correifolia and M. reitzii). Myrceugenia glaucescens flowers mainly in December and January and

matures its fruits in Februaryto April so it is an exception to the general rule. The different phenological rhythms sometimes serve to separate apparently closely related species and seem to have significance,at least sometimes, above the species level.

Myrceugenia

Table II Presence of Vessels with ScalariformPerforationPlates and Spiral Thickenings in Wood Samples Examined

Name 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20.

Luma apiculata M. acutiflora M. alpigena var. alpigena M. alpigena var. rufa M. campestris M. euosma M. fernandeziana M. glaucescens var. latior M. lanceolata M. miersiana M. miersiana M. mvrcioides var. acrophylla M. ovata var. acutata M. ovata var. gracilis M. oxysepala M. planipes M. pilotantha var. pilotantha M. pilotantha var. pilotantha M. reitzii M. rufescens

Collection Landrum 872 Pereira & Duarte 4522 Sucre 4655 Hatschbach 18158 Hatschbach 16471 Legrand 754 Pisano & Montaldo 1470 Hatschbach 7311 Rusbv 584 Reitz & Klein 8760 Hatschbach 10035 Hatschbach 14546 Hatschbach 3579 Hatschbach 1142 Hatschbach 11920 Landrum 947 Almeida 1265 Sucre 2462 Hatschbach 17182 Hatschbach 8247

At least some scalariform perforation plates present

Spiral thickenings present

yes no yes no no no no no no no no no yes yes no yes yes yes yes no

no weak yes yes no yes no weak weak no weak no yes no yes no no no no yes

Wood Anatomy There have been only cursory investigations of the wood anatomy of the AmericanMyrtaceae,but the little that is known indicates that Myrceugeniahas some characteristicswhich are rare in the rest of the family, namely vessels with scalariformperforationplates, tracheids,and vessels with spiralthickenings.The first two characteristicsare generallythoughtto be primitiveand the significance of the third is uncertain(Carlquist, 1975, pp. 194-195). Record and Hess (1943) in their treatment of the woods of the New World Myrtaceaereportedthat perforationplates were simple except in Luma apiculata (A. P. de Candolle) Burret [which they called Myrceugeniaapiculata (A. P. de Candolle) Niedenzu]; and that they had seen no spiral thickenings except in Myrceugenia schultzei and M. fernandeziana.

Metcalfe and Chalk (1950), in their anatomicaldescription of the family said that perforationplates were simple except for Myrceugenia, Myrtus communis L. and a species of Eugenia growing at Kew. They also reported spiral thickenings in some species of Eugenia and Myrceugenia. Little weight can be given this account because it is impossible to know how much was based on earlier publicationsand how much on their own work. It is possible that the species of "Eugenia" growing at Kew was Luma apiculata which is cultivated in Great Britain. Ingle and Dadswell (1953) found no scalariformperforation plates or spiral thickeningsin their extensive study of the woods of the Myrtaceaeof the south-

Flora Neotropica

western Pacific. These authors studied more than three hundred species of 32 genera plus two specimens of Myrtuscommunis, a mediterraneanspecies. Meyland and Butterfield(1975) have found scalariformperforationplates in

Neomyrtus pedunculata (Hook. f.) Allan. Ragonese (1976) found tracheids in Luma apiculata, Myrceugenia chrysocarpa, Amomyrtus luma (Mol.) Legrand et Kausel, Blepharocalyx gigantea Lillo

and B. tweediei (Hooker et Arnott) Berg. She did not find tracheids in the two species of Eugenia and one species each of Psidium and Campomanesiawhich she also examined. Dr. R. Schmid of the University of Californiaat Berkeley, informs me that the strand tracheids reported by Sastrapradjaand Lamoureux (1969) for Metrosideros are equivalentto those found by Ragonese. Schmid (pers. comm.) has recently found scalariformperforation plates in

Myrceugenia, Luma, Myrteola, Ugni and Neomyrtus. He has not been able to find them in Myrtus communis.

During my study of Myrceugenia I examined with the scanning electron microscope young wood specimens of 15 different species of Myrceugenia. Scalariformperforationplates were found in five species and spiral thickenings in nine species (Table II). Neither of these characteristics is uniformly present throughouta specimen, vessels also being found which have simple perforation plates and/or which have no spiral thickenings. Scalariformperforationplates were also found in the only specimen of Luma apiculata examined. The systematic value of scalariformperforationplates in Myrceugenia is not greatbecause of the variabilitywithinspecimens. It is always possible to overlook their presence. This and the difficulty of examining many specimens were the reasons for limitingthe survey. It is difficultfor the non-specialist to evaluate these rather sparse anatomical findings. As far as they go, the results of my observations and the accounts of previous workersseem to supportmy opinion(see ChapterIII), that Myrceugenia is one of the more primitivegenera of the Myrteae and that Luma and Blepharocalyx are fairly close relatives of Myrceugenia. ChromosomeNumbers Chromosomenumbershave usually proven to be systematicallyof little value in the Myrtaceaebecause of their invariability.In fact Atchison (1947)has stated, "In the Myrtaceae the lack of chromosome number variation is perhaps more pronounced than in any other dicotyledonous family of its size." N = 11 has been the common numberfound in almost every genus studied (Bolkhovskikhet al., 1969;Rye, 1979). Aneuploidshave been reportedfor some Australasiangenera (e.g. Actinodium, Darwinia, Homoranthus, Verticordia, Beaufortia, Thryp-

tomene) (Smith-White, 1948, 1950, 1954; Rye, 1979). And some polyploidy has been reported in a few genera (e.g. Baeckea, Callistemon, Eugenia, Leptosper-

mum, Psidium (Bolkhovskikhet al., 1969). Another problem encountered by anyone studying chromosomes in the Myrtaceae is their small size. Accordingto Atchison (1947)they are 1.2-2.5 microns long in Psidium and Eucalyptus. This makes exact counts difficultto obtain and thus prejudicemay creep in. Such prejudiceis indicatedperhapsby the fact that Harrison reported 2n = 28 in Eucalyptus citriodora Hooker and E. globulus La-

bill.; Sigiurareported 2n = 20 for the same species; Atchison reported 2n = 22 in E. citriodoraand McAulayet al. reportedn = 11 in E. globulus. These counts were all published or republished in Atchison (1947) and Bolkhovskikh et al. (1969). Although in all the observations which I have made n = 11 or 2n = 22

Myrceugenia Table III MaterialUsed for ObservingChromosomes Species Myrceugenia Myrceugenia Myrceugenia Myrceugenia Myrceugenia Mvrceugenia

bracteosa brevipedicellata euosma exsucca miersiana ovata var. gracilis

Collection

Type of material

Landrum 2817 Landrum 2830 Landrum 2719 Landrum 3084 Landrum 2978 Landrum 2637

young pollen young young young young

Landrum 2633 Landrum 3552 Landrum 3551

pollen mother cells root tips root tips

ovules mother cells ovules ovules ovules ovules and

megasporemothercell

Myrceugenia pilotantha var. major Myrcianthes fragrans Luma apiculata

appearto be probablenumbersI cannot say with certaintythat n does not equal 10, 12 or 14. In my judgmentonly polyploidsand significantlydifferentaneuploids can be of systematic value in the Myrtaceae. In this study several attempts were made to count chromosomes of species of Myrceugenia in preserved flower buds and with material grown in the greenhouse. Meiotic divisions were observed in pollen mother cells and megaspore mothercells, and mitotic divisions were observed in young ovules. All specimens of Myrceugenia examined were found to have n about equal to 11 or 2n about equal to 22. In addition,the root tips of specimens of Myrcianthesfragrans(Sw.) McVaughand Luma apiculata (A. P. de Candolle)Burret,grown at the University of MichiganBotanical Garden, were examined and also proved to have 2n about equal to 22. The species, collection and the type of materialexamined are listed in Table III. VI. SYSTEMATICTREATMENT Fruitingspecimens of Myrceugenia can be distinguishedby the myrcioid embryo from all other genera of the Myrteae except those of the Myrciinae. From other membersof the Myrciinae,the genus is distinguishedby: 4-merousflowers (occasionally 4-merous in Myrcia); few to several ovules in each of 2-4 locules; usually persistent bracteoles; and an inflorescence that is not a panicle. In Chile and adjacent Argentinafloweringspecimens of Myrceugenia are occasionally confused with the 4-merous genera Luma and Temu. Both the latter have bracteoles that are caducous at about anthesis, which distinguishes them from all the continental species of Myrceugenia. In eastern South America floweringspecimens of Myrceugeniaare most often confused with species of Eugenia. Eugenia usually has a bracteate shoot inflorescence whereas Myrceugenia sometimes has. In the Myrceugenia of eastern South America some peduncles are normallyvertically superimposed(except in M. ovata var. gracilis, M. smithii and M. hatschbachii); and the locule number

varies from two to four, some ovaries with three or four locules usually being present. In Eugenia superimposedpeduncles are absent (or at least rare)and the locule numberis essentially always two. In the preparationof this monographnumerous flowers were dissected and length and width measurementswere made of the bracteoles and calyx-lobes by cuttingthem away from the flowers andlayingthem on a scale. The measurements

Flora Neotropica

in the key and the descriptionsare of the bracteoles and the calyx-lobes as they are in the flowers, i.e. somewhat concave, not flattened. The peduncle width is a measurementat about 1 mm below the bracteoles. Separate keys for the geographic regions have not been constructed but the range of each species is indicatedin Table IV, so that one may quickly determine which species are found in a particularregion. Before attemptingto identify a specimen one may also find it useful to consult Table I, where the species are classified accordingto the location of dense hair cover. Many of the characterswhich are most useful in identificationare sometimes absent. Other normallyuseful charactersare variablein some species or may be interpretedin different ways. Thus many species appear more than once in this key. When one encounters a specimen that appears to be intermediatebetween two choices in the key, he should be able to identify it by following both leads. With little knowledge of Myrceugenia and with few specimens with which to compare, one may find identificationdifficult. Some experience with the genus and a good reference collection will do much to facilitate identification. Herbariumacronymsare those given in the Index Herbariorum(Holmgrenand Keuken, 1974). One herbariumnot listed in the index has also been consulted: that of the Instituto de Botanica of the UniversidadAustral, Valdivia, Chile for which the acronym VALD has been used. MYRCEUGENIABerg, Linnaea 27: 5 (in key). 1855; Linnaea 27: 131. 1856. Figs. 1, 5, 6, 7, 8. Mvrcia subgen. Myrceugenia(Berg) Kiaerskou,Enum. Myrt. bras. p. 115. 1893. Nothomyrcia Kausel, Lilloa 13: 148. 1948 (" 1947"). Type. Nothomyrciafernandeziana

(Hooker

et Arnott)Kausel [=Myrceugeniafernandeziana(Hooker et Arnott)Johow]. Mvrceugeniasect. Phanerocalvx Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 2 (in key). 1953;Darwiniana11(2):303. 1957. Mvrceugeniasubsect. Incanae Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 2 (in key). 1953;Darwiniana11(2):303. 1957. Mvrceugeniasubsect. Subulatae Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 3 (in key). 1953;Darwiniana11(2):303. 1957. Mvrceugeniasubsect. Aovatae Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 3 (in key). 1953;Darwiniana11(2):303. 1957. Mtrceugenia sect. Steganocalyx Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 2 (in key). 1953;Darwiniana11(2):303. 1957.Type. Myrceugeniamacrosepala (Burret)Legrandet Kausel [probablyequals Myrceugeniaoxysepala (Burret)Legrandet Kausel]. Myrceugeniasect. MicrobothrvsLegrand, Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 2 (in key). 1953;Darwiniana11(2):303. 1957. Myrceugeniasect. MacrobothrvsLegrand,Sellowia 13: 305. 1961.Invalidnamebecause no type was designated. Myrceugeniasubsect. UnifloraeKausel, Lilloa32: 351. 1966.Invalidname because no type was designated. Myrceugeniasubsect. Dichotomae Kausel, Lilloa 32: 354. 1966. Invalid name because no type was designated.

Trees up to ca. 10 m high or much branched shrubs less than 1 m high; hairs reddish-brown,yellowish or whitish, appressed or spreading,usually symmetrically or asymmetricallydibrachiate,less often simple, densely to sparsely covering (less often absent) twigs, leaves and flowers; leaves opposite (occasionally ternate or subopposite on some twigs), elliptic, ovate, lanceolate, obovate, oblanceolate or linear, thickly coriaceous to membranous, the midvein usually prominentbelow, usually impressed above, the marginaland lateral veins indistinct to prominent, the petiole usually channeled adaxially, rarely terete; inflorescence an axillary uniflorousdichasium, an axillary dichasiumwith 1-3 levels of bifurcation, or an axillary or terminal bracteate shoot (raceme); peduncles

Myrceugenia Table IV Distribution of Species of Myrceugenia

c Mo

0 0

c 0 ,

-E

.S O

M. correifolia M. obtusar

M. exsuccavar._____ M. lanceolata M. colchaguensis

M. pinifolia stern leptospermoides M. parvifolia M. ovata var. nannophylla M. ovata var. ovata M. chrvsocarpa M. planipes

0 oM0 bato 00

Masternspercies M. alpigena var. longifolia M. alpigena var. alpigena M. alpigena var. orufat M. rufescens M. campestris M. bracteosara M. myrcioides var. mvrcioides M. mvrcioides var. acrophylla M. miersiana M. ovata var. gracilis M. ovata var. acutata M. glaucescens var. latior M. scutellata M. seriatoramosa M. acutiflora M. pilotantha var. pilotantha M. franciscensis M. hoehnei M. brevipedicellata M. oxysepala M. hatschbachii M. kleinii

___

Flora Neotropica Table IV (continued)

73 o

c i .2

M.cucu3 ata

Eastern species M. M~ reitzii venosa M. glaucescens var. glaucescens M. cucullata M. euosma M. euosma M. venosa

M. M. pilotantha vat.major major pilotantha var. M. smithii M. mvrtoides M. leptocalyx

. ______

_________ ______

___

solitary or vertically superimposedin groups of 2-4(-5) in the axils of leaves and bracts; bracteoles ovate, lanceolate, linear, filiform, spatulate or orbicular,usually persistinguntil the fruit matures, rarely caducous at anthesis; calyx-lobes 4, broadly rounded to sharply acute, hemiorbicular,ovate or triangular,strongly concave to nearly flat, imbricate or less often valvate in the bud; hypanthium usually not prolonged beyond the summit of the ovary; stamens ca. 40 to over 500; petals 4, usually suborbicular;ovary 2-4-locular, the number of locules varyingfrom flower to flower; ovules normallyin two longitudinalrows, 2 to ca. 20 in each locule, the numbervaryingfrom locule to locule; fruit a fleshy berry, yellow, orange, reddish, or dark purple; seeds usually 1-5 per fruit, the testa membranous; embryo myrcioid, i.e. the cotyledons thin, membranous,folded into a bundle, the hypocotyl, horseshoe shaped, encirclingthe cotyledons. Type species. M. myrtoides Berg [lectotype, designatedby Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 3. 1953]. Myrceugenia-Main Key 1. Bracteoles scarious, caducous at anthesis;Mas a Tierra,JuanFernandez. 1. M. fernandeziana. 1. Bracteoles rarely scarious, not caducous at anthesis, usually persistinguntil the fruit matures. 2. Inflorescences mainly bracteate shoots, some dichasia and uniflorouspeduncles subtended by normalleaves also present. Key A. 2. Inflorescencesnever bracteateshoots, only dichasiaand uniflorouspedunclessubtended by normalleaves. 3. Some flowers borne in dichasia. Key B. 3. Flowers never borne in dichasia. 4. Bracteoles linearto filiform,4-9 times as long as wide. Key C. 4. Bracteoleslanceolate,ovate, orbicular,or spatulate,rarely4 times as long as wide. 5. Hairs all simple. Key D. 5. Hairs all dibrachiateor a mixtureof dibrachiateand simple. 6. Twigs and flowers yellowish lanate. Key E. 6. Twigs and flowers not yellowish lanate, mainlyreddish-brownpubescentor glabrous,the hairs usually appressedor spreading.

Myrceugenia

7. Leaves less than 2 cm long, densely pubescent beneath. Key F. 7. Leaves if less than 2 cm long, then not densely pubescentbeneath. 8. Pedunclesglabrousor with only scatteredhairs. Key G. 8. Pedunclespubescent. 9. Calyx-lobes triangular-ovate,concave throughoutinner surface, valvate, tightly closed in the bud, the closed bud a somewhatindented hemisphere. 31. M. myrcioides. 9. Calyx-lobesif triangular-ovate,then usually not concave throughout inner surface, imbricate, the closed bud not indented, often pointed. 10. Calyx-lobescoriaceousexcept for a thin membranousmargin. Key H. 10. Calyx-lobesmembranousto coriaceous,if coriaceous, without a thin membranousmargin. 11. Leaves 3-15 times as long as wide. Key I. 11. Leaves less than 3 times as long as wide. 12. Outer surface of the calyx-lobes and the hypanthium both densely pubescent, withouta sharpcontrastbetween them. 13. Calyx-lobesrounded;eastern and western South America. Key J. 13. Calyx-lobesacute; eastern South America. Key K. 12. Outer surface of calyx-lobes and the the hypanthium both glabrousor nearly so or the outer surfaceof the calyx-lobes notably less pubescent than the densely pubescent hypanthium. 14. Calyx-lobesacute. Key L. 14. Calyx-lobesrounded. Key M.

Key A 1. Hairs whitish, simple; midveinnot impressed;Mas a Tierra,JuanFernandez. 1. M. fernandeziana.

1. Hairs reddishbrownor yellowish brownor if whitishthen dibrachiate;midveinimpressed. 2. Pedunclesglabrousor nearly so. 3. Longest calyx-lobes about 1.7 mm long; peduncles about 0.3-0.5 mm wide at 1 mm below the bracteoles. 8d. M. ovata var. acutata. 3. Most calyx-lobes over 1.8 mm long, some normallyover 2 mm long; pedunclesabout 0.5-1.3 mm wide at 1 mm below the bracteoles. 20b. M. glaucescens var. latior. 2. Pedunclespubescent. 4. Bracteate shoots not over 1.5 cm long, terminatingin a flower; peduncles bearing 3. M. campestris. solitary flowers not exceeding 2 mm long; southernBrazil. 4. Bracteate shoots exceeding 2 cm long, terminatingin a flower or not; peduncles bearingsolitaryflowers exceeding 2 mm long. 5. Pubescence reddishbrown to yellowish brown, spreading,the hairs a mixtureof simple and asymmetricallydibrachiateor all simple. 6. Bracteoles 1-2 mm long, aboutV/ as long as the hypanthium;dichasianormally present; flowering mainly from Januaryto March; western South America. 5. M. exsucca.

6. Bracteoles 2-5 mm long, as long as or longer than the hypanthium;dichasia never present;floweringfrom Septemberto October;Brazil. 2. M. rufescens. 5. Pubescence whitish, appressed;hairs symmetricallydibrachiate. 7. Calyx-lobes3-6 mm long, 3.5-8 mm wide, densely pubescentwithout. 17. M. reitzii.

7. Calyx-lobes 1.8-3 mm long, 1.9-4 mm wide, sparsely pubescent to glabrous without. 8. Bracteateshoots terminatingin a flower;some petioles exceeding 1 cm long; 16. M. kleinii. floweringfrom August to September;Brazil. 8. Bracteate shoots not terminatingin a flower; petioles rarely exceeding 0.8 cm long; floweringmainlyfrom Decemberto March;western South America.

15. M. planipes.

Flora Neotropica

24 Key B

1. Twigs and flowers yellowish to greyish lanate. 34. M. myrtoides. 2. Calyx-lobesacute, 4-7 mm long; Rio Grandedo Sul and Uruguay. 4. M. schultzei. 2. Calyx-lobesrounded, 1-3 mm long; Mas Afuera, Juan Fernmndez. 1. Twigs and flowers not yellowish or greyish lanate. 3. Some petioles exceeding 1 cm long; pubescence whitish, appressed;most leaves ex16. M. kleinii. ceeding 5 cm long; Brazil. 3. Petioles not exceeding 0.8 cm long; pubescence usually reddish-brown,not appressed, or if whitishand appressedthen the leaves not exceeding 5 cm long. 4. Peduncles bearing dichasia not exceeding 6 mm long, the flowers of the dichasia 3. M. campestris. sessile or the pedicels up to 3 mm long; Brazil. 4. Peduncles bearingdichasiaexceeding 6 mm long, the flowers of the dichasia sessile or with definitepedicels over 3 mm long. 5. Bracteoles0.3-1 mm wide; hypanthium2-5 mm long; western South America. 6. Leaves 1-2.5 times as long as wide; dichasiausuallywith more than 3 flowers; 5. M. exsucca. twigs not reddish;bracteolesusually less than 0.5 mm wide. 6. Leaves 3-15 times as long as wide; dichasiausually with no more than 3 flow6. M. lanceolata. ers; twigs reddish;bracteolesusually 0.5-1 mm wide. 5. Bracteoles 1-3 mm wide; hypanthium1-2 mm long; eastern South America. 7. Calyx-lobesroundedto blunt, about as wide as long; hairs on twigs all dibrachiate, usually appressed;leaves usually widest above the middle;peduncles 26b. M. alpigena var. rufa. usually 2-4 togetherin the axils of the leaves. 7. Calyx-lobesacute, longer than broad; hairs on the twigs a mixtureof simple and dibrachiate,spreading;leaves usually widest near the middle;peduncles 28. M. euosma. usually 1-2 in the axils of the leaves.

Key C 1. Bracteolesleaf-like,petiolate;leaves 1-2.5 cm long (shade leaves sometimeslonger),ovate 10. M. obtusa. to elliptic, generallyobtuse, the midveinnever excurrent;centralChile. 1. Bracteoles not leaf-like, withouta noticeablepetiole, usually widest at the base; leaves 28 cm long, variouslyshaped, the apex generallyacute, the midveinoften excurrent;eastern South America. 2. Petals pubescentwithout;matureleaves pubescentbeneath,often over 6 cm long, often with prominentlateralveins; floweringmainlyfrom Januaryto May. 3. Leaves with prominentlateraland marginalveins; pubescence yellowish to goldenbrown, the hairs all simple or nearly so, those on the lower surface of the leaves 33. M. miersiana. somewhatcurled and erect. 3. Leaves with lateral and marginalveins not prominent;pubescence reddish-brown, the hairs usually a mixtureof simple and asymmetricallydibrachiate,those on the lower surfaceof the leaves straightto slightlycurled, usually not erect. 32. M. pilotantha.

2. Petals glabrousor nearlyso without;matureleaves glabrousor nearlyso beneath,rarely over 6 cm long (except in M. campestris), the lateral veins not prominent;flowering from Marchto October. 4. Flowering mainly from August to October; calyx-lobes usually blunt to rounded, mostly under 3 mm long, mostly less than 1.5 times as long as wide; not becoming reflexedas they mature. 5. Flowers sessile or nearly so, borne on maturetwigs, the fertile nodes sometimes

3. M. campestris.

leafless.

5. Flowers with peduncles4-12 mm long, borne on young leafy twigs. 2. M. rufescens. 4. Flowering(as far as known) from Marchto July; calyx-lobes (at least two) sharply acute, 3-4 mm long, usually 1.5-2 times as long as wide, becomingreflexedas they mature.

35. M. hoehnei.

Key D 1. Hairs whitish; midveinnot impressed;Mas a Tierra,Juan Fernandez. 1. M. fernandeziana. 1. Hairs yellowish to reddish-brown;midveinimpressed;eastern South America. 2. Petals pubescent without;floweringmainlyfrom Januaryto May; some maturecalyxlobes usuallyreflexed;matureleaves pubescentbeneath,often over 6 cm long, the lateral veins often prominent. 33. M. miersiana.

Myrceugenia 2. Petals glabrousor nearly so without;floweringmainlyfrom August to October;mature calyx-lobes curlinginwards,never reflexed;leaves pubescentbeneathwhen young, glabrescent at maturity,rarelyover 6 cm long, the lateralveins not prominent.

2. M. rufescens.

Key E 1. Calyx-lobes4-7 mmlong; bracteoles4.5-8 mmlong; most hairsasymmetricallydibrachiate; 34. M. mvrtoides. Rio Grandedo Sul and Uruguay. 1. Calyx-lobes 1-4 mm long; bracteoles 1-4.5 mm long; hairs asymmetricallydibrachiateto symmetricallydibrachiateor simple. 2. Hairs all simple or nearly so; lateralveins often prominent;petiole terete or nearly so; 33. M. miersiana. petals pubescentwithout;bracteoles0.5-1.2 mm wide. 2. Hairsall dibrachiateor a mixtureof aboutV/ dibrachiateand /2 simple(in the lattercase the bracteoles 1.5-3 mm wide); lateralveins rarelyprominent;petiole channeled;petals glabrousor nearly so without. 3. Bracteoles0.5-1 mm wide; western South Americaand Juan Fernandez. 4. Leaves mainly 1-2 cm long (shade leaves occasionallylonger), often verticillate, the apex broadly rounded; calyx-lobes not greenish, coriaceous, more or less 24. M. colchaguensis. acute; mainlandChile. 4. Leaves 1.5-5(-6) cm long, some normallywell over 2 cm long, not verticillate,the apex often acute or acuminate;calyx-lobes greenish, roundedor blunt, membra4. M. schultzei. nous; Mas Afuera, Juan Fernandez. 3. Bracteoles 1-3 mm wide; eastern South America. 5. Hairsall symmetricallydibrachiate;leaves usuallywidest above the middle;calyxlobes roundedto blunt;from Rio Grandedo Sul to MinasGerais. 26b. M. alpigena var. rufa.

5. Hairs a mixtureof simple and dibrachiate;leaves usually widest near the middle; one pairor all the calyx-lobesusuallyacute;from Camposdo Jordaoin Sao Paulo northto MinasGerais and Rio de Janeiro. 27. M. hracteosa.

Key F 1. Leaves with the midvein indistinguishableor nearly so above; peduncles 1-4 mm long; 7. M. rufa. calyx-lobes rounded;centralChile. 1. Leaves with the midvein distinguishableabove; peduncles (3-)4-20 mm long; calyx-lobes generallyacute; eastern South America. 2. Hairson the twigs all dibrachiate,appressed;hypanthium(1.5-)2-3 mmlong; bracteoles 26a. M. alpigena var. alpigena. mainly 2-4 times as long as wide. 2. Hairson the twigs a mixtureof dibrachiateand simple, spreading;hypanthium1-1.8 mm 28. M. euosma. long; bracteoles mainly 1-1.8 times as long as wide.

Key G 1. Bracteoles more or less orbicular,obtuse, about 1-2 mm long, somewhatwider than long; Brazil.

21. M. scutellata.

1. Bracteoles not orbicular,usually ovate to lanceolate, longer than wide. 2. Twigs glabrousor inconspicuouslypubescent, the hairs (if any) small, appressed;peduncles often in pairs. 3. Longest calyx-lobes about 1.7 mm long; peduncles about 0.3-0.5 mm wide at 1 mm below the bracteoles;Brazil. 8d. M. ovata var. acutata. 3. Most calyx-lobesover 1.8 mm long, some normallyover 2 mm long; pedunclesabout 0.5-1.3 mm wide at 1 mm below the bracteoles. 4. Bracteoles mainly2-2.7 times as long as wide; floweringmainlyin Februaryand March, the fruits maturingthe next season mainly in December and January; leaves 1.5-5 cm long, often all under2.5 cm long, the apex usuallyobtuse; calyxlobes never densely pubescentwithin;western South America. 19. M. chrysocarpa. 4. Bracteolesmainly 1.2-2 times as long as wide; floweringmainlyin Decemberand January,the fruitsmaturingthe same season in February,March,andApril;leaves 2.5-8 cm long, the apex usually acute; calyx-lobes sometimesdensely pubescent 20. M. glaucescens. within;eastern South America. 2. Twigs densely pubescent, at least when young;peduncles almost always solitary. 5. Leaves mainly3-7 times as long as wide.

Flora Neotropica 6. Bracteoles more than 1 mm long, about as long as the calyx-lobes; leaves coriaceous, the base cuneate; twigs pubescentwhen young, becomingglabrescentbefore the first bark exfoliates; Brazil. 13. M. hatschbachii. 6. Bracteoles less than 1 mm long, much shorterthan the calyx-lobes; leaves subcoriaceousto membranous,the base roundedto acute; twigs remainingpubescent 11. M. parvifolia. until the first barkexfoliates; Chile. 5. Leaves mainly 1.5-3 times as long as wide. 7. Calyx-lobesabout 1.5-2 mm long; bracteolesabout 1/2 as long as the calyx-lobes; 11. M. parvifolia.

Chile.

7. Calyx-lobesmainlyless than 1.5 mmlong;bracteolesabout2 as long as the calyx8c. M. ovata var. gracilis.

lobes; Brazil.

Key H 1. Stamens 250 to over 500; leaves 4-16 cm long; calyx-lobes (2.5-)4-6.5 mm long; twigs conspicuouslycovered with hairs, not soon glabrescent. 31b. M. myrcioidesvar. acrophylla. 1. Stamens70-240; leaves 2.5-6 cm long; calyx-lobes 1.5-4 mmlong;twigs glabrousto densely covered with hairs when young, soon glabrescent. 2. Hairs whitish; peduncles usually not exceeding 0.5 cm long; leaves membranousto 22. M. cucullata. subcoriaceous,the upper surfacedull, the apex sharplyacuminate. 2. Hairs reddish-brown;peduncles mainly 0.5-1 cm long; leaves coriaceous, the upper 23. M. seriatoramosa. surfacelustrous, the apex usually bluntlyacuminate.

Key I 1. Outer surface of calyx-lobes and the hypanthiumboth densely pubescent, with no sharp contrastbetween them. 2. Bracteoles 4-7 mm long, the apex acuminate;peduncles rarely exceeding 3 mm long; 36. M. brevipedicellata. Camposdo Jordao, Sao Paulo. 2. Bracteoles 1.5-4 mm long, the apex not acuminate;peduncles usually exceeding 3 mm long. 3. Leaves glabrousor nearly so beneath. 4. Twigs glabrousor inconspicuouslycovered with whitish appressedhairs; calyxlobes often with a small horn-likeextension near the apex; Brazil. 22. M. cucullata. 4. Twigs densely and conspicuouslypubescent,at least when young;hairssometimes spreading;calyx-lobes withouta small horn-likeextension near apex. 5. Leaves 5-8 cm long; bracteoles 1-1.7 times as long as wide; Goias, Brazil. 26c. M. alpigena var. longifolia.

5. Leaves 1-5 cm long;bracteoles2-4 times as long as wide; not growingin Goias. 6. Leaves oblanceolateto linear, often verticillate;twigs reddish;calyx-lobes rounded; Chile.

6. M. lanceolata.

6. Leaves narrowlyelliptic to lanceolate, not verticillate;twigs not reddish; calyx-lobes usually acute, less often rounded;Brazil. 26a. M. alpigena var. alpigena.

3. Leaves, at least when young, densely pubescentbeneath. 7. Petioles terete or nearlyso; petals pubescent;bracteoles2-5 times as long as wide; hairs on the twigs all simple or a mixtureof simple and dibrachiate. 8. Leaves with prominentlateral and marginalveins; pubescence yellowish to golden-brown,the hairs all simple or nearly so, those on the lower surface of the leaves somewhatcurledand erect. 33. M. miersiana. 8. Leaves with lateral and marginalveins not prominent;pubescence reddishbrown, the hairs usually a mixtureof simple and asymmetricallydibrachiate, those on the lower surfaceof the leaves straightto slightlycurled, usually not erect.

32. M. pilotantha.

7. Petioles channeled;petals glabrousor with a few scatteredhairs; bracteoles 1-5 times as long as wide, if over 2 times as long as wide then the hairs on the twigs all dibrachiate. 9. Hairs on the twigs all dibrachiate,bracteoles lanceolate, 2-4 times as long as wide; hypanthiumabout as long as or longerthan the calyx-lobes.

26a. M. alpigena var. alpigena.

9. Hairs on the twigs a mixtureof simple and dibrachiate;bracteoles 1-1.8 times as long as wide; hypanthiumusually shorterthan the calyx-lobes.

Myrceugenia

10. Peduncles mostly 2-4 together in a leaf axil, mostly under 0.5 cm long; leaves 3-6 cm long, the marginoften stronglyrevolute; leaves and twigs 27. M. bracteosa. tomentose; Camposdo Jordao, Sio Paulo northward. 10. Peduncles mostly solitary or two together in a leaf axil, mostly over 0.5 cm long; leaves 1-3(-5) cm long, the marginsnot stronglyrevolute;leaves and twigs sericeous, especially on the lower leaf surface; Paranasouthward.

28. M. euosma.

1. Outer surface of the calyx-lobes noticeablyless pubescentthan the densely pubescenthypanthium. 11. Calyx-lobes0.5-0.8 times as long as wide, most exceeding 3 mm wide; leaves narrowly elliptic, often exceeding 5.5 cm long and 2 cm wide; hairs whitish, appressed;Chile. 15. M. planipes.

11. Calyx-lobes0.8-1.7 times as long as wide, rarelyexceeding3 mmwide; leaves narrowly elliptic, linearor oblanceolate,rarelyexceeding 5.5 cm long and 2 cm wide; hairsoften reddish-brownand spreading. 12. Leaves oblanceolateto linear; twigs reddish;hypanthium1.5-2 times as long as 6. M. lanceolata. wide, usually slightly contractedjust below the calyx-lobes;Chile. 12. Leaves elliptic to linear; twigs not reddish;hypanthiumusually about as wide as or wider than long, not contractedjust below the calyx-lobes. 13. Leaves 1-2 cm wide; twigs glabrousor inconspicuouslypubescent, the hairs small and appressed;peduncles often in pairs; calyx-lobes often with a small horn-likeextension near apex; Brazil. 22. M. cucullata. 13. Leaves 0.2-1 cm wide; twigs densely pubescent, the hairs usually spreading; pedunclesalmost always solitary;calyx-lobes withouta smallhorn-likeextension near the apex. 14. Hypanthium1.5-2 mmlong;leaves stronglydiscolorous,darkgreenabove, light green below; stamens about 100-150;calyx-lobes about as long as or shorterthan the hypanthium;Chile. 9. M. pinifolia. 14. Hypanthiumabout 1 mm long or shorter;leaves more or less concolorous, light grey-green;stamensabout 60-100; calyx-lobesabout 1.5 times as long as the hypanthium. 15. Leaf base abruptlyobtuse; bracteoles blunt; twig pubescence short, appressed; Chile.

12. M. leptospermoides.

15. Leaf base cuneate, the leaf graduallytaperingfrom the middle;bracteoles acute; twig pubescence spreading;Brazil. 14. M. smithii.

Key J 1. Twigs densely pubescent. 2. Hairs all dibrachiate. 26. M. alpigena. 2. Hairs a mixtureof simple and dibrachiate. 33. M. miersiana. 1. Twigs glabrousor inconspicuouslypubescent. 3. Pubescence whitish; peduncles usually not exceeding 0.5 cm long; leaves membranous to subcoriaceous,the upper surfacedull, the apex sharplyacuminate. 22. M. cucullata. 3. Pubescence reddish-brown;peduncles mainly 0.5/1 cm long; leaves coriaceous, the 23. M. seriatoramosa. upper surfacelustrous, the apex usually bluntlyacuminate.

Key K 1. Bracteoles 4-7 mm long, lanceolate; calyx-lobes acute, triangular,3-5 mm long, 1.6-2.8 times as long as wide; peduncles rarelyexceeding 3 mm long; petals glabrous. 2. Leaves mainly 3-5 times as long as wide, the apex acute, the midvein often slightly excurrent;calyx-lobes pubescent within except sometimes at the base; bracteole apex 36. M. brevipedicellata. acuminate;known only from Camposdo Jordao, Sao Paulo. 2. Leaves mainly 2-2.8 times as long as wide, the apex usually rounded or obtuse, the midvein not excurrent;calyx-lobes glabrous or nearly so within except at the apex; bracteole apex acute to obtuse; mainly Santa Catarinaand Rio Grandedo Sul, rare in Paranaand Sao Paulo. 37. M. oxysepala. 1. Bracteoles various sizes, in most species not exceeding 4 mm long, if longer than 4 mm long the calyx-lobes not over 1.6 times as long as broad;pedunclesoften over 3 mm long; petals often pubescent. 3. Twigs glabrousor only sparsely and inconspicuouslycovered with minutehairs.

Flora Neotropica 4. Hairs whitish; peduncles usually less than 0.5 cm long; leaves membranousto sub22. M. cucullata. coriaceous, the upper surfacedull, the apex sharplyacuminate. 4. Hairs reddish-brown;peduncles mainly 0.5-4.5 cm long; leaves subcoriaceousto coriaceous, the upper surface often lustrous, the apex sometimes bluntlyacuminate or acute. 5. Peduncles 1-4.5 cm long; leaf apex usually sharplyacute, sometimes mucronate; leaves densely pubescent when young. 25. M. franciscensis. 5. Peduncles0.4-1 cm long; leaf apex usuallybluntlyacute or acuminate;leaves not 23. M. seriatoramosa. densely pubescent when young. 3. Twigs densely pubescent. 6. Hairs on the twigs dibrachiate,generallysymmetrical. 7. Peduncles(1-)2-4.5 cm long; leaves glabrescentor nearly so at maturity. 8. Leaves often exceeding4 cm long; hypanthiumaboutas long as or shorterthan 25. M. franciscensis. calyx-lobes; bracteolesca. 1.5-2 times as long as wide. 8. Leaves rarelyexceeding 4 cm long; hypanthiumusuallylonger than the calyx26a. M. alpigena var. alpigena. lobes; bracteolesca. 2-4 times as long as wide. 7. Peduncles0.3-1(-2.2) cm long; leaves densely pubescentbeneathat maturity. 9. Leaves obovate to oblanceolate;calyx-lobes blunt, about as long as broad; peduncles usually 2-4 in a row in the leaf axils; petals glabrousor nearly so without.

26b. M. alpigena var. rufa.

9. Leaves elliptic, sometimes narrowlyso; calyx-lobes acute, triangular,usually longer than broad;pedunclesusually 1-2 in a row in the leaf axils. 10. Petals densely pubescentwithout,calyx-lobes3-5.5 mm long; leaves often exceeding 1.5 cm wide.

29. M. acutiflora.

10. Petals glabrousor nearly so without;calyx-lobes 1.6-3.2 mm long; leaves 26a. M. alpigena var. alpigena. usually not exceeding 1.5 cm wide. 6. Hairs on the twigs all simple or a mixtureof simple and dibrachiate,the lattermainly asymmetrical. 11. Leaves 1-4 cm long, the lower surfaceglabrousor very sparselypubescent, the lower epidermismore or less shiny and wrinkledwhen dry; knownonly fromthe Sao Francisco de Paula regionof Rio Grandedo Sul. 38. M. leptocalvx. 11. Leaves more than 4 cm long or densely pubescentbeneath;lower epidermisnot shiny, not wrinkledwhen dry. 12. Marginaland lateralveins prominent,ca. 0.2 mm in diam. 13. Lower leaf surface densely covered (at least when young) with simple yellowish to golden-brownhairs, these slightly curled and more or less 33. M. miersiana. erect; petiole unchanneledor only slightly sulcate. 13. Lower leaf surface sparsely covered with dibrachiatehairs, these not 30. M. venosa. erect; petiole channeled. 12. Marginaland lateralveins not as prominent,less than 0.2 mm in diam. 14. Petals densely pubescent without;petiole unchanneledor only slightly sulcate. 15. Pubescenceyellowishto golden-brown,the hairsall simpleor nearly so, those on the lower surface of the leaves somewhat curled and erect; calyx-lobesrarelyacute; leaves rarelyaveragingover 3 times as long as wide; (exceptions to the last two charactersbeing found in Rio Grandedo Sul). 33. M. miersiana. 15. Pubescencereddish-brown,the hairsusuallya mixtureof simpleand asymmetricallydibrachiate,those on the lower surfaceof the leaves straightto slightlycurled, usually not erect; calyx-lobes sometimes acute; leaves sometimesaveraging3-4 times as long as wide. 32. M. pilotantha.

14. Petals glabrousor with only scatteredhairs without;petiole channeled. 16. Pedunclesmostly 2-4 per leaf axil, mostly under0.5 cm long; leaves 3-6 cm long, more or less tomentose beneath; Campos do Jordao northward.

27. M. bracteosa.

16. Pedunclesmostly 1-2 per leaf axil, mostly over 0.5 cm long; leaves 1-3(-5) cm long, sericeous beneath;Paranasouthward. 28. M. euosma.

Key L 1. Twigs densely covered with reddish-browndibrachiatehairs. 26a. M. alpigena var. alpigena. 1. Twigs glabrousto sparselycovered with reddish-brownor whitish hairs.

Myrceugenia 2. Pubescence appressed,whitish, often farinose on the hypanthium;peduncleslackingor up to 5(-7) mm long, 0.5-1 mm below the bracteoles, densely pubescent, not subtended by bracts;hypanthiumusually 1.5-2 mmlong; flowerbud often longerthanthe peduncle.

22. M. cucullata.

2. Pubescence reddish-brown,more or less spreading;peduncles 3-18 mm long, 0.3-0.6 mm wide at 1 mm below the bracteoles;glabrousto sparselypubescent, often subtended by bracts; hypanthiumusually about 1 mm long; flower bud rarely longer than the peduncle.

8d. M. ovata var. acutata.

Key M 1. Pedunclesdensely covered with reddish-brownhairs. 2. Leaves mainly 1-3 cm long; twigs densely pubescent; hairs a mixture of simple and asymmetricallydibrachiate. 3. Bracteoles 0.5-2 mm long, lanceolate to ovate, clasping the hypanthium,not leaf8. M. ovata. like; eastern and western South America. 3. Bracteoles 2-6 mm long, elliptic, linear, oblong or spatulate,not claspingthe hypan10. M. obtusa. thium, leaf-like, petiolate;Chile. 2. Leaves often exceeding 3 cm long; twigs glabrous or sparsely pubescent to densely pubescent;hairs appressed,all symmetricallydibrachiate. 4. Twigs and lower leaf surfaces sparselypubescentto glabrous;leaf apex bluntlyacuminate; Brazil.

23. M. seriatoramosa.

4. Twigs and the lower surfaces of at least young leaves densely pubescent;leaf apex rarelyacuminate,usually rounded. 5. Bracteoles 3-8 mm long; leaf marginsstronglyrevolute; Chile. 18. M. correifolia. 5. Bracteoles 1.4-2.5 mm long; leaf marginsnot stronglyrevolute;Brazil. 26b. M. alpigena var. rufa.

1. Peduncles sparselypubescentor if densely pubescent, the pubescence whitish. 6. Calyx-lobes 1-1.7 mm long. 7. Pubescence appressed, whitish, often farinose on the hypanthium;peduncle lacking or up to 5(-7) mm long, 0.5-1 mm wide at 1 mm below the bracteoles, densely pubescent, not subtendedby bracts;hypanthiumusually 1.5-2 mm long; flower bud often longer than the peduncle. 22. M. cucullata. 7. Pubescencereddish-brown,more or less spreading;peduncles3-18 mm long, 0.3-0.6 mm wide at 1 mm below the bracteoles, glabrousto sparsely pubescent, often subtendedby bracts;hypanthiumusuallyabout 1 mm long; flowerbud rarelylongerthan the peduncle.

8d. M. ovata var. acutata.

6. Most calyx-lobes over 1.8 mm long, some normallyexceeding 2 mm long. 8. Some petioles exceeding I cm long; some leaves usually over 7 cm long and 3 cm 16. M. kleinii. wide; Brazil. 8. Petioles rarelyexceeding 0.8 cm long; leaves usuallyneithermore than7 cm long nor 3 cm wide. 9. Calyx-lobesusually not over 2 mm wide, often with a small horn-likeextension near the apex; peduncles lacking or up to 5(-7) mm long, densely covered with whitishappressedhairs;hypanthiumusually not over 2 mm long, obconic; leaves 22. M. cucullata. rarelyover 5 cm long; Brazil. 9. Calyx-lobes often over 2 mm wide, never with a small horn-likeextension near the apex; pedunclesusuallyall over 5 mm long, glabrousor pubescent, if as short as 5 mm and also densely covered with whitishappressedhairs(as in M. planipes occasionally), then the hypanthiumhemispherical,usually over 2 mm long and some leaves longerthan 5 cm. 10. Hypanthium(1.5-)2-4 mmlong, densely covered with minuteappressed,whitish hairs;peduncles ca. 1 mm wide, usually densely glandular;leaves almost all acuminateat both apex and base, mainly3-8 cm long; fruit maturingfrom November to December,almost a year after flowering;western South Amer15. M. planipes. ica, generallyat altitudesbelow 700 m. 10. Hypanthium1-2 mm long, glabrousto densely pubescent, the hairs whitish, yellowishor reddish-brown;peduncles0.5-1 mm wide, withoutglandsor glandular;leaves often roundedor bluntat apex or base, often all under3 cm long; eastern and western South America. 11. Bracteoles mainly 2-2.7 times as long as wide; floweringmainlyin February and March, the fruits maturingthe next season in December and January;leaves 1.5-5 cm long, often all under 2.5 cm long, the apex

Flora Neotropica usually obtuse; calyx-lobes never densely pubescent within; western 19. M. chrysocarpa. South America, mainlyat 700-1000 m. 11. Bracteoles mainly 1.2-2 times as long as wide; floweringmainly in December and January,the fruits maturingthe same season in February, March and April; leaves 2.5-8 cm long, the apex acute, acuminate or rounded;calyx-lobes sometimesdensely pubescentwithin;eastern South America.

20. M. glaucescens.

1. Myrceugeniafernandeziana(Hooker et Arnott) Johow, Flora de Juan Fernandez 94. 1896. Fig. 9.

MyrtusfernandezianaHooker et Arnott, Bot. Misc. 3: 316. 1833. Eugenia lumilla Philippi, Bot. Zeitung (Berlin) 14: 643. 1856. Type. PresumablyPhilippi s.n. "JuanFernandez."(holotype, SGO, n.v.; Field Museumneg. 31577of probableisotype at W, photo at MICH). Eugeniafernandeziana(Hooker et Arnott)Barneoudin Gay, Fl. chil. 2: 392. 1847. Myrceugenialuma Berg, Linnaea30: 671. 1860.(Illegitimatenamebecause Myrtusfernandeziana Hooker et Arnottis cited in synonymy.) Lumafernandeziana(Hookeret Arnott)Burret,Notizbl. Bot. Gart.Berlin-Dahlem15: 526. 1941. Nothomvrciafernandeziana(Hooker et Arnott)Kausel, Lilloa 13: 148. 1948("1947").

Tree up to 25 m high; hairs simple, usually spreading,whitish; twigs moderately to sparsely pubescent when young, glabrescent with age; leaves very sparsely pubescent to glabrous, ovate to lanceolate, 2.8-7.3 cm long, 1.2-2.3 cm wide, 2-4 times as long as wide; apex acute to acuminate,the tip usually blunt; base slightly cordate to narrowly rounded; petiole 2-5 mm long, moderately pubescent to glabrouswhen young losing any pubescence with age, often rough with cracks and wrinkles, barely channeled, reddish-brownto blackish; midvein about flush with the blade to slightlyraised, prominent,with longitudinalwrinkles above and below; lateral veins prominent,up to about 40 pairs visible, the reticulate secondaryveins numerous;marginalveins equallinglateralsin prominence, archingslightly between them; blades light yellow-greenor grey-greento darker grey-greenabove (rarelydryingreddish-brown),the same color or slightlylighter below, coriaceous;flowers mainly aggregatedtogether in bracteate shoots of 412, the bracteate shoots 1-3 cm long, either solitary or seriate at the nodes, terminatingin a vegetative bud or flower; peduncles solitary or in pairs in the axils of bracts (perhaps also occasionally in the axils of leaves), 2-11 mm long, ca. 0.5 mm wide, densely to sparselypubescent; bracts and bracteoles ovate, 12.5 mm long, ca. 1 mm wide, quite membranous,deciduous at anthesis or before, sparsely pubescent to glabrous within and without, some bracts serving as bud scales; calyx-lobes ovate, strongly concave, more or less acute, 1.6-4 mm long, 1.6-3.5 mm wide, 1-1.4 times as long as wide, membranousto slightly fleshy, sparsely pubescent without, sparsely to densely pubescent within; hypanthium obconic, densely pubescent, 1.5-3 mm long; disk 2-3 mm across, moderatelyto densely pubescent, sometimes becoming slightlyconvex duringmaturation;style densely to sparsely pubescent, 6-9 mm long; stamens 40-80, 5-10 mm long; anthers 0.5-0.8 mm long when dry; petals more or less orbicular to slightly elongate, 2-4 mm in diam.; ovary 2-3-locular; ovules (1-)3-7 per locule; fruit orange when fresh, darkenedwhen dry, few seeded. Type. Douglas s.n. "Juan Fernandez." (hereby designated as lectotype, E-GL); Scouler s.n. (paratype, n.v.).

Specimensexamined.CHILE. Isla de Mas a Tierra(RobinsonCrusoe):Plazueladel Yunque,May 1938, Webers.n. (H), Nov 1940, Webers.n. (H), elev. ca. 250 m, 20 Sep 1955, Sparre & Planella 140 (H), 250 m und hoher, Oct 1934,Bock 37 (F, GH, MO, NY, US); Pico Central,ca. 400 m, 20 Oct 1955, Sparre & Planella 198 (H); faldeos del Yunqueentre la Plazoletay el Camote, ca. 300 m, 1116 Aug 1942,Pisano & Montaldo1444(UC); faldeos de los cerros de BahiaCumberland,100-250m,

Myrceugenia

11-16 Aug 1942, Pisano & Montaldo 1421 (H, UC); cerros entre Centinelay el Pangal, ca. 300 m, 11-16 Aug 1942,Pisano & Montaldo1470(H, UC, US); Portezuelode Villagra,9 Dec 1965, Solbrig 3809 (LP, MICH,NY, UC); CerroSalsipuedes,2010ft, 12 Dec 1965,Meyer9527 (MICH,UC), 1380 ft, 10 Dec 1965, Meyer 9509 (MICH, UC); Cord6nSalsipuedes, 21 Apr 1917, Skottsberg75 (NY); withoutspecificlocality, 10-14 Jan 1901, Hastings248 (NY, UC, US), Nov 1875,Moselevs.n. (GH); Nov 1941, Webers.n. (CONC, H, SI).

Berg took up the name Myrceugenialuma for this species, citing Myrtusluma Bertero as a basionym. According to Johow (1896) Myrtus luma Bertero is a nomen nudum and I have no reason to doubt that he is correct. Myrtus luma Molina (Saggio sulla st. Nat. Chili: 173. 1782) is Amomyrtus luma (Molina) Kausel, accordingto Kausel(1947).Gunckel(1972)has madethe new combination Nothomyrcia maxima (Molina) Gunckel, based on Myrtus maxima Molina,

publishedin the above cited work. Gunckel apparentlybelieves that N. maxima should replace N. fernandeziana (Hooker et Arnott) Kausel. I believe that it is impossible to identify M. maxima as any of the known Chilean Myrtaceae because of the vague protologue and in particularbecause it is supposed to have alternateleaves, which none of the ChileanMyrtaceaehave. The firstconfirmable, legitimatename for the species here underconsiderationwas Myrtusfernandezi-

ana Hooker et Arnott, so Mvrceugenia fernandeziana is the name that should be used in Myrceugenia.

Myrceugeniafernandeziana is one of the most distinctive species in the genus but it has no characteristicthat cannot be found in some other species. Therefore it has not been retained as a separate genus as was proposed by Kausel. If Kausel had been familiarwith the eastern membersof the genus he mighthave recognizedthe similaritybetween M.fernandeziana and the two Brazilianspecies M. rufescens and M. campestris. These all have bracteate shoot inflorescences (i.e. racemes), flowers of about equal size and they share a relatively low ovule and stamen number. Mvrceugenia fernandeziana has, and M. rufescens usually has, simple hairs. Myrceugenia rufescens has linear bracteoles, M. fernandeziana

has ovate bracteoles and those of M. campestris range between these two types. In M. campestris the bracteateshoot terminatesin a flower, in M. fernandeziana it sometimes does and in M. rufescens it terminates in a vegetative bud. Both M. fernandeziana and M. rufescens have a disk which occasionally becomes

swollen during fruit development and also have some calyx-lobes which are both acute and concave throughoutthe entire inner surface, a combinationnot commonly found in the calyx-lobes of Myrceugenia. Mvrceugenia rufescens

and M. campestris both have linear bracts and flower early in the spring. All three species normallygrow in approximatelythe same habitat, i.e. foggy, subtropical forests near the ocean. Numerical analyses indicate that these three species are a closely related group although all the charactersjust mentioned were not includedbecause they were not applicableto all the species in the genus (Landrum, 1981). Mvrceugeniafernandeziana is easily distinguishablefrom all the other species in the genus by its scarious bracteoles that are caducous at about anthesis and its large leaves with the midveins not impressed and somewhat wrinkled. It flowers from August to December and the fruits probably mature in July and August. It is the dominanttree on Mas a Tierra Island and according to Johow (1896) the wood of this species is very hard and resistant and has been used "desde tiempos inmemoriales"for buildingboats. He also states that the fruits can be used as a spice in cooking in the same way as "pimienta de Jamaica," probably referringto Pimenta, the allspice. A map showing the distributionof

M. fernandeziana is given in Fig. 9.

Flora Neotropica

2. Myrceugeniarufescens (A. P. de Candolle) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 8. 1953. Figs. 8A-B, 9, 27A. Eugenia rufescens A. P. de Candolle,Prodr.3: 279. 1828. Eugenia poeppigiana Berg, in Mart. Fl. Bras. 14(1): 232. 1857. Type. Sellow s.n. "Habitat in

fruticetisrepublicaeMontevideo,floretSeptembriet Octobri"(holotype, B, apparentlylost; Field Museumneg. 31602of possible isotype at W, photo at MICH). Eugenia planiramea Berg, in Mart. Fl. Bras. 14(1): 234. 1857. Type. Pohl 1013, "Habitatad vicum Bordo do Campo in prov. Minarum"(holotype, W; Field Museum neg. 31598 of holotype, photo at MICH). Luma rufescens (A. P. de Candolle)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 528. 1941. Lumaplaniramea (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 527. 1941. Lumapoeppigiana (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 533. 1941. Mvrceugeniarufescens var. alegrensis Legrand,in Legrandet Klein, Flora Ilustr. Catarinense [Mirt]:433. 1970. Type. Reitz & Klein 4780, Santa Catarina,Campo Alegre, Morro do Iquererim,elev. 1400m, 5 Sep 1957(holotype, MVM;isotypes, HBR, US). Mvrceugeniajonssonii Kausel, Lilloa 33: 106. 1971. Type. Jonsson 998a. Brasil, Parana,Itaperussu, elev. 880 m, 27 Sep 1914(holotype, S, n.v.; isotype, MICH).

Shrub or small tree 1-5 m high; hairs all simple or a mixture of simple and dibrachiate,reddish-brownto yellowish; twigs densely pubescent when young, glabrescentwith age, the barkremainingsmooth or becomingrough with cracks, grey to reddish-brown;leaves densely pubescent when young, losing some or all pubescence with age, generally more pubescent beneath than above, elliptic to obovate or oblanceolate, 2.6-5.3 cm long, 0.9-2.3 cm wide, 2.5-3.5 times as long as wide; apex rounded to acute, the midvein often slightly excurrent; base cuneate or acuminate;petiole 2-5 mm long, 1-1.5 mm thick, densely pubescent when young, glabrescent with age, slightly channeled; midvein impressed for entire length or only proximallyabove, prominent,slightly sulcate below; lateral veins indistinguishableor up to about 10 pairs faintly visible; marginal veins equalling laterals in prominence; blades light to dark grey-green above, light yellow-green to reddish-brownbeneath, coriaceous; flowers often grouped together in bracteate shoots of up to ca. 12 flowers, the bracteate shoots up to ca. 3.5 cm long, axillary or terminal, terminatingin a vegetative bud; peduncles solitary or in pairs, 4-12 mm long, 0.5-1 mm wide, densely pubescent; bracts linear, 3-4 mm long, 0.5-1 mm wide, densely pubescent; bracteoles linear, 3-4 mm long, 0.5-0.7 mm wide, densely pubescent throughoutor glabrousproximally within; calyx-lobes ovate, more or less concave, 2.2-3.4 mm long, 2-3.2 mm wide, 1-1.5 times as long as wide, submembranous,densely to sparselypubescent without, densely to sparselypubescentdistallyto glabrousor nearlyso proximally within; hypanthiumobconic, densely pubescent, 1.5-2.3 mm long; disk ca. 2 mm in diam., sparsely pubescent, sometimes becoming enlarged and convex after flowering;style 4-6 mm long, very sparsely pubescent to glabrous;stamens 80120, 3-8 mm long; anthers 0.4-0.7 mm long when dry; petals suborbicular,3-5 mm in diam.; ovary 2-4-locular; ovules 7-12(-14) per locule; fruit subglobose, 3-5 mm in diam., blackish-purple,pubescent; seeds usually 1-2 per fruit, ca. 4 mm long. Type. Martius s.n., "in Brasiliae campis prov. S. Pauli" (holotype, M, n.v.; Field Museum neg. 19996of holotype at M, photo at MICH). Specimens examined. BRAZIL. Minas Gerais: Conceigao de Ibitipoca, ca. 1200 m, 3 Oct 1970, Sucre et al. 7273 (RB); prope S. Antonio, Ponte Nova, Aug 1896, Robello s.n. (R). Sao Paulo: Sao Paulo, 1 Sep 1914, Pickel 5374 (US); Sao Paulo, Villa CerqueiraCezar, 14 Sep 1918, Hoehne s.n. (MBM, SP); Sao Paulo, nativa no JardimBotanico, 28 Sep 1931,Hoehne s.n. (NY, SP); Sao Paulo, Ypiranga,Nov 1913, Luederwaldts.n. (SP), Aug 1907, Luederwaldts.n. (SP); Avenida Paulista, 16 Sep 1906, Usteris.n. (SP); Sao Bernardo,26 Oct 1913,Brades.n. (MBM, SP); Sao Paulo, Vila Ema,

Myrceugenia

Oct 1953, Brade 21290 (HB, MVM, RB); Itarare,Campos de Sao Pedro, Serra de Bom Sucesso, Fazenda Ventania, ca. 1000 m, 10 Dec 1966, J. Mattos 14888 (SP), 20 Oct 1966, J. Mattos 14027 (SP). Parana:Mun. Bocaiuva do Sul, Tunas, 11 Sep 1961, Hatschbach 8247 (HB, HBR, MBM, MICH, MVM);Mun. Guarapuava,Guara,18Jan 1968,Hatschbach 18309(C, MBM, MICH,MVM). Santa Catarina:Mun. CampoAlegre, Morrodo Iquererim,1500m, 5 Sep 1957, Reitz & Klein 4788 (HBR, MICH, US), 900 m, 18 Oct 1957, Reitz & Klein5191 (HBR, US).

Myrceugenia rufescens is probably most closely related to M. fernandeziana

and M. campestris and is compared to them in Key A. It is more likely to be confused with M. hoehnei and M. pilotantha and is comparedwith both in Key C and in the discussions of those species. It might also be confused with M. miersiana, which can be distinguishedby its prominentlateraland marginalveins, unchanneledpetiole, densely pubescent petals and floweringperiod. Myrceugeniarufescens seems to be found mainlyin the misty forests along the eastern edge of the planalto and in the Serra Paranapiacaba.A map of its distribution is shown in Fig. 9. It flowers mainly in September and October and the fruits probably maturein December and January. 3. Myrceugeniacampestris(A. P. de Candolle) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 12. 1953. Figs. 9, 27B. Eugenia campestris A. P. de Candolle,Prodr.3: 274. 1828. Eugenia distans Berg, in Mart. Fl. Bras. 14(1):263. 1857.Type. Sellow s.n., "Habitatin prov. S. Pauli" (holotype, B, apparently lost; Field Museum neg. 36948 of an apparent isotype at

P, photo at MICH). Luma campestris (A. P. de Candolle)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 529. 1941. Luma distans (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 529. 1941.

Myrceugenia distans (Berg) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 12. 1953. Myrceugenia campestris var. distans (Berg) Legrand, Flora Ilustr. Catarinense [Mirt]: 447. 1970.

Small tree or shrub 2-4 m high; hairs reddish-brown,a mixtureof dibrachiate and simple; twigs densely to sparsely pubescent when young, glabrescent with age, the bark smooth whitish-grey;leaves generally elliptic, less often ovate or obovate, 2.5-11(-16.5) cm long, 1-4.5 cm wide, 1.5-3.5 times as long as wide, glabrous to sparsely puberulent above, glabrous to sparsely pubescent below; apex usually acute or acuminate,less often rounded, the midvein usually excurrent; base rounded,cuneate or acuminate;petiole densely to sparselypubescent, channeled, 1-4 mm long, 1-2 mm thick; midveinimpressed, narrowabove, prominent below; lateral veins indistinctor up to about 17pairseasily visible; marginal veins equallinglateralsor less distinct; blades dull darkgrey-greenabove, lighter yellow green or with a reddish cast below, subcoriaceousto membranous;inflorescence uniflorous, a dichasiumor a compact bracteate shoot with up to about 9 flowers, arising in the leaf axils or at leafless nodes of the year-old twigs; peduncles lacking to 2 mm long, about 1 mm wide, densely pubescent; bracteoles linear to ovate, sparsely to densely pubescent without, less densely pubescent to glabrous within, membranousto subcoriaceous; calyx-lobes ovate to oblong, slightly concave, membranous,sparsely pubescent without, sparsely pubescent to glabrouswithin; hypanthiumobconic, densely pubescent, 1.5-2.2 mm long; disk 1.5-3 mm across, sparsely pubescent to glabrous; style 5-7 mm long, sparsely puberulent to glabrous; stamens about 40-70, 3-8 mm long; anthers 0.4-0.5 mm long when dry; petals white, more or less orbicular,about 2-3 mm in diam.; ovary usually 2-locular, less often 3-locular;ovules (2-)4-8 per locule; fruit pubescent, globose, 4-6 mm in diameter;seeds 3-4 mm long. Type. Martius s.n., "in Brasiliae campis prov. Minarum," "v. s. in herb.

Flora Neotropica

Mart." (holotype M, n.v.; Field Museum neg. 19922of holotype at M, photo at MICH). Specimens examined. BRAZIL. Sao Paulo: Morro do Jaragua,7 Sep 1925, Hoehne s.n. (SP); Sales6polis, Estacao Biol6gica de Boraceia, 12 Sep 1961, J. Mattos 9102 (MBM, SP), 16 Apr 1964, J. Mattos 11770 (SP); Paranapiacaba,Estagao Biol6gicaAlto da Serra, 9 Aug 1961,J. Mattos 9069 (HB, MBM, SP). Parana:Mun. CampinaGrandedo Sul, Serrada Lapinha,4 Aug 1963, Hatschbach 10183(HB, MBM, MICH, MVM);Mun. CampinaGrandedo Sul, Sito do Belizario, 1000m, 18 May 1967, Hatschbach 16423 (MBM, MICH, MO, NY, UC), 4 Oct 1967, Hatschbach 17303 (MBM,

MICH,MVM, NY); Mun. CampinaGrandedo Sul, Caminhoao CerroVerde, 21 May 1967,Hatschbach 16471(C, CTES, F, HB, MBM, MICH,MVM, NY); Mun. Paranagua,Rio Guaraguassil,29 Jul 1951, Hatschbach 2433 (MBM, MVM); Mun. Paranagua,Sertao do Indaial, 10 m, 8 Nov 1965, Hatschbach 12742 (F, HBR, MBM, MICH, MVM, US); Mun. Guaratuba,Rio da Divisa, 5 m, 17 Aug 1970, Hatschbach24660 (C, HB, MBM, NY, UC); Mun. Guaraquefaba,Serrinha,9 Aug 1967, Hatschbach 16893 (MICH); Mun. Piraquara, Manancias da Serra (ca. 25?30'S, 49?W), ca. 1000 m, 31

Oct 1977, Landrum2274, 2275 (MBM, MICH); Mun. Morretes, Via Graciosa, Grota Funda (ca. 25020'S,48?50'W),13 Dec 1977, Landrum2873, 2874, 2880 (MBM, MICH). Santa Catarina:Mun. Brusque, Sao Pedro, 31 Aug. 1950, Klein 112 (H, HBR, PACA); Mun. Itajai, Cunhas, 10 m, 29 Sep 1955, Klein 1600 (H, HBR, MVM, NY, US); Mun. Palhofa, Campodo Massiambi,5 m, 5 Nov 1953, Reitz & Klein 1292 (H, HBR, MVM, US); Mun. Sao Franciscodo Sul, MinaVelha, Garuva, 10 m, 5 Oct 1957, Reitz & Klein 4986 (H, HBR, MVM). Rio Grandedo Sul: Sao Leopoldo, Portao, 20 Jul 1949, Rambo 42654 (MVM).

Myrceugeniacampestris is easily distinguishedfrom all other species of Myrceugenia by its compact inflorescences of glomerules of short bracteate shoots or dichasia. When the inflorescence is not fully developed, the combinationof linearbracts, short peduncles, roundedcalyx-lobes and leaves withoutprominent veins distinguishes it from all the other species. It blooms mainly in the spring months of August and Septemberand the fruits mature shortly thereafter. Myrceugenia campestris typically grows in the misty forests along the eastern edge of the planalto and in the coastal lowlands. A map of its distributionis shown in Fig. 9. 4. Myrceugenia schultzei Johow, Flora de Juan Fernandez: 96. 1896. Figs. 7B, 9. Luma schultzei (Johow) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 526. 1941.

Tree up to 12 m high; hairs yellow to yellowish-brown,almost entirely symmetrically dibrachiate;twigs densely lanate when young, glabrescent with age, the young twig bark greyish to yellowish-white, sometimes tinged with reddishbrown, persisting as strips; leaves elliptic, ovate, obovate, lanceolate or oblanceolate, 1.5-5(-6) cm long, 0.5-3 cm wide, 1.4-2.8 times as long as wide, densely lanate below when young, densely lanate to glabrousabove, glabrescenton both surfaces with age, the smaller, narrowerleaves on flower bearingbranches;apex acuminate, acute to rounded; base acuminate, cuneate, acute or obtuse; petiole channeled, 1-4 mm long, 0.5-1.5 mm thick, densely lanate when young, sometimes glabrescent; midvein shallowly impressed proximally, to not at all impressed above, prominentbelow; lateral veins indistinct or up to about 10 pairs faintly visible; marginal veins about as prominentas laterals or less so; blades coriaceous, grey-green, yellow-green or tinged with reddish-brownabove, the same color or somewhat lighter below; peduncles uniflorousor bearing a three (rarely more) flowered dichasium, flattened, 0.5-2 cm long, ca. 0.5 cm wide, densely lanate (rarelyglabrous),solitaryor up to 3 in a row in the axils of leaves, the terminalflower of the dichasiumsessile, the lateralflowers with stalks similar to but shorterthan the uniflorouspeduncles, these stalks occasionally in pairs in the axils of the bracteoles of the terminalflower; bracteoles ovate to lanceolate,

Myrceugenia

1-2 mm long, 0.8-1 mm wide, 1.2-2.3 times as long as wide, submembranous, densely to sparselycovered with hairs, rarelyglabrouswithout,glabrousor nearly so within, clasping the hypanthium;calyx-lobes ovate-oblong, 1-3 mm long, 12.2 mm wide, 0.8-1.5 times as long as wide, sparsely covered with hairs (rarely glabrous)without, sparsely covered with hairsto glabrouswithin, concave; petals glabrous or nearly so, suborbicular,2-3.5 mm in diam.; hypanthium obconic, 1.5-2.5 mm long, densely lanate; disk 2-3 mm across, sparsely covered with hairs to glabrous; stamens 90-140, 4-7 mm long; anthers orbicularto oblong, 0.3-0.5 mm long; style 4-6 mm long, glabrous or nearly so; ovary 2-4-locular; ovules 6-12 per locule; fruit unknown. Type. "Germain, Ag. Guajardo, Johow," "Masafuera," (lectotype, hereby designated, Johow s.n. "Masafuera, 31.XII.91," SGO-069757). Specimens examined. CHILE. Isla de Mas Afuera (Alejandro Selkirk): Apr 1941, Weber s.n. (CONC, H), 28 Dec 1891, Johow s.n. (SGO-069721), 1 Jan 1892, Johow s.n. (SGO-069824); 1700 ft, 30 Jan 1935, Chapin 70 (GH, NY), Chapin 1071 (NY, US); Quebrada Blindado, 30 Nov 1965, Solbrig 3711 (GH, UC); Quebrada de la Mata Maqui on trial to Correspondence Camp, 1300 ft, 3 Dec 1965, Meyer 9442 (MICH); Quebrada del Mono, 12 Feb 1917, Skottsberg s.n. (C, NY, US), 1 Dec 1965, Solbrig 3730 (GH); Quebrada de las Vacas, 21 Jan 1955, Skottsberg 134 (NY).

This species, which is endemic to the island of Mas Afuera, may be most closely related to Myrceugenia exsucca of the continent. Myrceugenia schultzei

can be distinguishedfrom the other species in the genus by its combinationof dichasial inflorescence and yellowish, symmetricallydibrachiatehairs. It seems to flower from Januaryto April and probablythe fruits mature sometime during the winter months when the island is uninhabited.A map of its distributionis shown in Fig. 9. 5. Myrceugeniaexsucca(A. P. de Candolle)Berg, Linnaea 30: 671. 1861. Figs. 8C-D, 9, 27C. Eugenia exsucca A. P. de Candolle, Prodromus 3: 278. 1828. ?Eugenia temu Hooker et Arnott, Bot. Beech. Voy.: 56. 1832. Type. Cruckshanks s.n., Mathews s.n. "Hab. Valparaiso," (syntypes E-GL?, n.v.). Eugenia multiflora Hooker et Arnott, Bot. Misc. 3: 322. 1833, not Eugenia multiflora Lam., Encyc. 3: 302. 1789 nor Eugenia multiflora Cambessedes, in Saint-Hilaire, Flora Brasiliae meridionalis 2: 361. 1833. Type. Bridges 320, "Quillota" (holotype, E-GL; Bridges specimen without number possible isotype at K; Field Museum neg. 31593 of isotype at W, photo at MICH). ?Luma temu (Hooker et Arnott) Gray, U.S. Expl. Exped., Phan.: 539. 1854. Mvrceugenia lechleriana Berg, Linnaea 27: 133. 1856. Type. Lechler 563, "Habitat in Chile prope urban Valdivia" (holotype, W; isotypes, GH, RB). Myrceugenia camphorata Berg, Linnaea 27: 134. 1856. Type. Poeppig s.n., "Chile" (holotype W; Field Museum neg. 31398 of holotype, photo at MICH). Eugenia exsucca [var.] a peruviana Berg, Linnaea 27: 255. 1856. (Inadmissible name to be replaced by E. exsucca var. exsucca because E. exsucca A. P. de Candolle is cited in synonymy). Eugenia exsucca [var.] p patagua Berg, Linnaea 27: 256. 1856. Type. Philippi 136, "Chili" (holotype, W). Eugenia pitra Berg, Linnaea 27: 264. 1856. (New name for Eugenia multiflora Hooker et Arnott.) ?Eugenia exsucca [var.] y temu (Hooker et Arnott) Berg, Linnaea 27: 256. 1856. ?Eugenia exsucca [var.] 6 apiculata Berg, Linnaea 27: 257. 1856. Type. Chamisso 56 "Chili" (holotype, B, n.v., apparently lost). Eugenia corralensis Philippi, Linnaea 33: 72. 1864. Type. Krause s.n. "Prope Corral" (holotype, SGO, n.v.; isotype, MICH; Field Museum neg. 36859 of isotype at P, photo MICH; Field Museum neg. 19928 of isotype at M, photo at MICH). Myrceugenia multiflora (Hooker et Arnott) Kausel, Rev. Mirt. Chile: 2. 1940; Revista Argent. Agron. 9: 63. 1942. Luma exsucca (A. P. de Candolle) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 525. 1941. Luma pitra (Berg) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 525. 1941.

Flora Neotropica

Luma corralensis(Philippi)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 525. 1941.

Tree up to ca. 12 m high; pubescence reddish-brownto yellowish, the hairs simple or asymmetricallydibrachiate;twigs densely pubescent when young, the hairs more or less spreading,graduallydeciduous or persisting until the young bark exfoliates; leaves suborbicular, elliptic, elliptic-obovate or elliptic-ovate (1-)2-7.5(-12.5)

cm long, (0.8-)1.5-3.5(-5)

cm wide, 1.2-2.4 times as long as

wide, sparsely puberulentto glabrous above, moderatelyto sparsely pubescent below when young, glabrescentwith age; apex and base acute to roundedor less often acuminate;petiole unchanneledor weakly channeled, 1.5-6 mm long, 1-2 mm thick, sparsely to densely pubescent; midvein impressedfor entire length or only proximally above, prominentbelow, often slightly excurrent; lateral veins indistinct or up to about 20 pairs visible; marginal veins equalling laterals in prominenceor less prominent;blades coriaceous to subcoriaceous, dark or light grey-green to reddish-brownabove, lighter yellow-green to reddish-tanbelow, usually strongly discolorous, the surfaces dull; inflorescence a solitary flower, a dichasiumor less often a bracteateshoot, the multiflorousinflorescences3-10 cm long, bearingup to 10flowers;peduncles solitaryor up to 3 in a row in the axils of leaves and bracts, flattened, 0.5-1 mm wide, densely pubescent; bracteoles lanceolate, 1-2 mm long, 0.3-0.5 mm wide, 2-5 times as long as wide, sparsely to densely pubescent withinand without, subcoriaceous;calyx-lobes ovate to ovateoblong, concave, 1.6-2.8 mm long, 1.7-2.7 mm wide, 0.7-1.2 times as long as wide, subcoriaceous, sparsely to densely pubescent within and without, strongly reflexedafter flowering;hypanthiumobconic to subglobose, 2-4 mm long, densely pubescent; disk 2.5-3 mm across, densely to sparsely pubescent, slightly recessed in the center; stamens 170-275, 3-10 mm long; anthers 0.3-0.5 mm long

when dry; petals glabrousto sparsely pubescent, suborbicular,2-4 mm in diam., white to tan when dry; style 5-8 mm long, sparselypubescent; ovary 2-4-locular; ovules 7-13 per locule; fruit globose, about 6-8 mm in diam., yellow-brownto orange-brownwhen dry; seeds 2-9, orbicularto oblong, 3-5 mm long. Type. Dombey s.n., "in Peruvia," "v. s. comm. a cl. Jussieu" (holotype, P, n.v.; Field Museum neg. 7941 of apparentholotype, photo at MICH).

CHILE. Coquimbo:FundoPalo ColoradoII, Quebradadel Buitre,zona litoral, 14Oct 1948,Kausel 2640 (CONC, F). Aconcagua:Zapallar,Jan 1899, Johow s.n. (SGO). Valparaiso:Quintero,26 Feb 1941, Looser 4360 (F, SGO), 10 Jan 1952, Poulsen s.n. (C); El Criquet, 18 Sep 1932, Looser 2574 (F); Vina del Mar, Quebradadel Tranque,18 Aug 1940, Kausel 784 (CONC, F, LP); Limache,Caj6n del rio San Pedro, 26 Feb 1932,Garaventa2905 (F); Limache,Cerrosde - ?, 11Feb 1931,Garaventa 2145 (GH); Caminode Placillaa Quintay,Feb 1958, Kausel 4341 (F, H), Kausel4342 (F); San Juan (Placilla/Quintay),4 Feb 1941, Kausel 906 (F), Kausel 907 (F, HBR, SGO);Algarrobo,14 Dec 1941, Kausel 997 (F), Kausel 996 (SGO);Algarrobo,Quebradadel Hotel Pacifico, Feb 1957, Kausel 4333 (F, H), Feb 1958, Kausel 4350 (H); Algarrobo,QuebradaLos Claveles, 4 Mar 1952, Boelcke 6503 (MO), 6504 (CTES, MO); Algarrobo, ca. 33 km north of San Antonio (ca. 33?22'S, 71?40'W), Quebrada

del Hotel Pacifico, 10-30 m, 26 Mar 1978, Landrum3442 (MICH, SGO); Las Dichas, 7 Mar 1968, Kausel5397, 5397a (H); Lagunilla,Cumings102 (F). Santiago:Malvillaca. 10km east of San Antonio (ca. 33?35'S,71?31'W),ca. 100m, 27 Mar 1978,Landrum3446, 3451 (both MICH, SGO). Colchagua: Antivero, zona cordilleranade San Fernando,27 Nov 1938, Kausel458 (F); Aguas Buenas, ca. 800 m, 21 Apr 1971, Landrum& Donoso s.n. (EIF). Curic6:Los Quenes, 19 Jan 1941, Kausel 846 (F, H), 18 Jan 1941, Kausel 850 (CONC, F, H, HBR); zona cordillerana,PotreroGrande,Ranchillo,12/ 20 Dec 1940, Bernath s.n. (F); San Pedro de Alcantara, 17 Feb 1946, Kausel 1840 (CONC, H). Linares: Laguna Amargo, ca. 50 km east of Parral, ca. 720 m, 21 Feb 1972, Landrum920, 921 (MICH). Maule: Coronel ca. 19 km southwest of Cauquenes, ca. 200 m, 24 Feb 1972, Donoso, Landrum& Gajardo s.n. (EIF); Constituci6n,20 Mar 1940, Kausel 703 (H, HBR). Concepci6n: Laraquete,24-25 Mar 1940,Kausel 736, 737 (F, H), Kausel 735 (H); Escuadron,25 Mar 1940,Kausel 760 (F); ca. 8 km east of Concepci6n(ca. 36?50'S,73?0'W),ca. 100 m, 27 Jan 1978, Landrum3084 (MICH, VALD); Parque Hualpen ca. 15 km west of Concepci6n, near sea level, 29 Jan 1978, Landrum3110 (MICH, VALD); Concepci6n, Mertens s.n. (GH). Arauco: Lago Lleu-lleu, 24 Feb

Myrceugenia

1939,Kausel 606 (H); Isla Mocha, La Calera,Oct 1958, KunkelM328 (H). Bio-Bio: Laja, caminode Santa Barbaraa Rio Huequecura(37?42'S,71?49'W),290 m, 12 Mar 1976, Marticorena,Quezada&

Rodriguez 894 A (CONC). Malleco: Puren, 27 Feb 1939, Kausel 620 (F), Kausel 619 (H), Feb 1939, Kausel 561 (F), Kausel 567 (F), Kausel 557 (F), Kausel 562 (H), Sep 1939, Kroll s.n. (F, H, LP);

23 Mar 1954,Sparre& Constance10861(CONC, UC). Cautin:Lautaro, camino Lautaro-Curacautin, 21 Feb 1964, Kausel 4802 (H); Quepe, Fundo Fin-Fin, Pitranco (38048'S,72?40'W),1 Jun 1970, Roivainen3218 (H); Nupangue, 20-23 Mar 1939, Bernath s.n. (F, H); Loncoche, 15-23 Mar 1939, Bernaths.n. (F, H); Tolten, Mar 1935,Friedrich1826 (CONC, F); Huiscapi, Feb 1941, Saelzer s.n. (F); Temuco, 15 Jan 1906, Sargent s.n. (A); Almagro, 60 m, 21 Feb 1943, Gunckel 14247 (LIL). Valdivia:San Jose de la Mariquina,ca. 50 m, Oct 1925, Hollermayers.n. (GH, MO, NY, SI, US); Panquipulli,ca. 200 m, Mar 1925, Hollermayers.n. (MO, NY, US); about 10 km north of Valdivia (ca. 39?45'S,73?15'W),ca. 100 m, 24 Feb 1978, Landrum3273 (MICH, VALD); Rinihue, 19 Feb 1941, Kausel 943 (CONC, H); Hosteria de Pirihueico, 600 m, 13-15 Mar 1948, Kausel 5461 (F); Corral,1862,Philippis.n. (LP, MICH);Niebla, Jan 1923,Gunckel371 (GH);Valdivia,1896,Buchtien s.n. (GH, US), Apr 1904,Buchtiens.n. (GH, US), 1888,Philippi727 (US). Osorno:Lago Rupanco, 8 Feb 1972,Cantino40 (GH). Llanquihue:Ensenada,49 km east of PuertoVarason Lago Llanquihue (ca. 41?15S, 72?30'W),ca. 50 m, 22 Feb 1978, Landrum3232, 3242 (MICH, VALD), 23 Mar 1939, Morrison17601 (GH, MO, UC). Chilo6:camino a Chepu ca. 25 km south of Ancud, 8 May 1972,Landrum946 (MICH);Putemun,cerca de 15 km al norte de Castro, 4 Mar 1942, Munoz2761 (H); Castro, 10 Jan 1924, Barros s.n. (H); Nelcon, a 10 km al sur de Castro, camino a Chonchi, 4 Mar 1942, Munoz 2756 (F); Huildad, near sea level, 8 Feb 1978, Landrum3163 (MICH, VALD); Yaldad,ca. 10 km west of Quell6n(ca. 43?15'S,73?45'W),1-5 m, 7-8 Feb 1978,Landrum3124, 3137 (MICH, VALD). Provinceunknown:near Rio Colorado,2500 ft, 29 Jan 1902, Hastings 474 (NY). ARGENTINA.Neuqu6n:Pucaraat Lago Lacar (40?S),725 m, 14 Dec 1955,Bocher, Hejerting& Rahn 1675 (C), 24 Feb 1953, Schajouskoy s.n. (LIL); Hua Hum, 14 Dec 1937, Kalela 1449 (H), 21 Jan 1947, Barba 1756 (F), Barba 1778 (MO), 2 Feb 1948, Dawson & Schwabe 2337 (MICH), 3 Mar

1949, Milano s.n. (MICH);Quetrihu6,costa Lago Nahuel Huapi, 29 Mar 1942, Diem 326 (F, H); Quetrihue,costa Lago Patagua, 20 Feb 1940, Diem 330B (H); Peninsula Quetrihue,Lago Nahuel Huapi (ca. 40?50'S,71?30'W),ca. 800 m, 27 Feb-2 Mar 1978, Landrum3288, 3311, 3316, 3321 (LP, MICH);PeninsulaQuetrihu6,Los Arrayanes, 1 Mar 1978, Landrum3300, 3301 (LP, MICH);Lago Nahuel Huapi, Isla Victoria, Puerto Anchorena, 780 m, 30 Mar 1934, Spegazzini 302-(No. 9) (MICH).Rio Negro: PuertoPanuelo, Lago Nahuel Huapi(ca. 41?S,71?30'W),3 Mar 1978,Landrum 3334, 3336 (both LP, MICH);Lago Nahuel Huapi, Llao Llao, 8 Apr 1934, Lycinger [?] 999 (NY); Lago Nahuel Huapi, ColoniaSuiza, 780 m, 12 Mar 1945,Barba376 (UC); Lago Nahuel Huapi,Bahia Lopez, 17 Mar 1939, Maldonado21 (LP, NY), 10 Mar 1943,Reuthells.n. (LIL); Lago Mascardi,8 Dec 1937, Kalela 1336 (H); El Bols6n, 12 Feb 1946, Boelcke 2096 (CTES). Chubut:Lago Puelo, 10 Nov 1945, T. Meyer 9304 (F), Feb 1942, Perez Moreau 573 (LIL); camino del Lago Puelo a El Bols6n, 7 Feb 1944, Nicora 3950 (CTES, F); Lago Puelo, desembocaduradel Epuy6n, 13 Feb 1940, Perez Moreau s.n. (LIL); Epuy6n, Feb 1945, Martinez Crovetto 3225 (LIL).

No type has been found for Eugenia temu Hooker et Arnott but another collection, Cuming 100, which Hooker and Arnott cite as Eugenia temu (Bot. Misc. 3: 321. 1833.) has been seen and is M. exsucca. A type specimen for Eugenia exsucca [var.] 6 apiculata has not been found but it is likely that it is also a synonym because Berg's specific concept was usually narrow. The hybridization which takes place between Myrceugenia exsucca and M. lanceolata and M. ovata var. nannophylla will be discussed in the section on hybridization. Myrceugenia exsucca is one of the most common constituents of the lowland myrtaceous forests of southwestern South America. A map of its distribution is shown in Fig. 9. It often grows in swamps, around lakes, and along streams with its roots entirely submerged. According to Gay (1847) this species is used in baths for the pains of rheumatism. Myrceugenia exsucca flowers from January to March and the fruits seem to mature from September to November. 6. Myrceugenia lanceolata (Jussieu ex Jaume Saint-Hilaire) Kausel, Lilloa 13: 135. 1948 ("1947"). Figs. 7C, 7F-G, 9, 27D. Myrtuslanceolata Jussieu ex Jaume Saint-Hillaire,in Duhamel, Trait6arbr. arbust., ed. 2, 1: 208. 1800-1803.

Flora Neotropica Eugenia dombevana A. P. de Candolle,Prodr.3: 276. 1828. New name for Myrtuslanceolata Jussieu ex Jaume Saint-Hilaire,proposed because of the existence of Eugenia lanceolata Lam. Mvrtusgudilla Colla, Mem. Accad. Sci. Torino 37: 66. (preprintsMay 1833)late 1834. Type. Berteros.n. "PenualasProvinciaeValparadisiacae"(holotype, TO, n.v.; photo of apparent holotype from TO seen). EugeniastenophyllaHookeret Arnott,Bot. Misc. 3: 322. 1833.Type. Cuming98 (herebychosen as lectotype, E-GL; isolectotype GH); Cruckshankss.n., "near Valparaiso"(paratype,EGL); Bridges319, 'near Valparaiso"(paratype,E-GL);Gilliss.n., "nearTalca" (paratypes, E-GL, GH); Menzies s.n., "Chili" (paratype,n.v.); Cuming97 (paratypes,E-GL, K). Eugenia gudilla (Colla)Barneoud,in Gay, Fl. chil. 2: 396. 1847. Luma stenophvlla(Hooker et Arnott)Gray, U.S. Expl. Exped. Phan.: 540. 1854. Eugenia stenophylla [var.] a angustifolia Hooker ex Berg, Linnaea 27: 254. 1856. (Inadmissible name to be replaced by Eugenia stenophylla var. stenophylla.) Eugenia stenophylla [var.] f latifolia Hooker ex Berg, Linnaea 27: 254. 1856. Type. Cuming 97

"v. in hb. Vindob. et Mart." (isosyntypes, E-GL, K). Myrceugeniastenophylla(Hooker et Arnott)Berg, Linnaea30: 670. 1861. Luma dombeyana(A. P. de Candolle)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 525. 1941.

Shrubup to 3 m high; hairs reddish-brownto whitish, a mixtureof simple and dibrachiate;twigs reddish when young, later brown or grey, densely to sparsely pubescent when young, soon glabrescent,the hairs spreading,mainly simple and asymmetricallydibrachiate;leaves oblanceolateto linear, (1.5-)2-5 cm long, 0.21.5(-2) cm wide, (2.5-)3-15 times as long as wide, glabrousor with a few scattered appressed dibrachiate hairs below; apex acute, acuminate, obtuse or rarely rounded; base acute or cuneate, normallytaperingfrom above the middle of the leaf; petiole unchanneledor weakly channeled, sparsely pubescent to glabrous, 1-4(-5) mm long, 0.5-1 mm thick; midvein shallowly impressedfor nearly entire length above, moderatelyprominentbelow; lateral veins indistinctor up to ca. 10 pairs of sharply ascending veins faintly visible; marginal veins equallingor less prominentthan the laterals; blades coriaceous, darkgrey-greento reddish-brown above, lightergrey-green,yellow-greenor tan below when dry, normallystrongly discolorous, the surfaces dull; peduncles uniflorousor bearing a 3-flowereddichasium (rarely with more than three flowers), flattened, 0.4-3.5 cm long, 0.5-1 mm wide, densely to sparsely pubescent, solitary or in pairs (rarely triplets) in the leaf axils, the terminalflower of the dichasiumusually without a stalk beyond the bracteoles; bracteoles of solitary flowers and lateral flowers in dichasia lanceolate, 1.5-3 mm long, 0.5-1 mm wide, 2.5-4 times as long as wide, subcoriaceous, sparsely to densely pubescent without, sparsely pubescent to glabrous within, clasping the hypanthium,the bracteoles of central flowers of dichasia similarbut larger,up to 15 mm long; calyx-lobes ovate to oblong, 1.8-4 mm long, 1.5-3 mm wide, 1-1.7 times as long as wide, submembranous,sparselypubescent distally to glabrous within, sparsely to moderatelypubescent without, concave, becoming stronglyreflexedafterflowering;petals glabrousto sparselypubescent, suborbicular,2-4 mm in diam., white to tan when dry; hypanthiummore or less obconic, contractedbelow the calyx-lobes, 2-5 mm long, densely pubescent; disk sparsely to densely pubescent, 2-3 mm across, recessed in the center; stamens 100-220, 3-7 mm long; anthers 0.3-0.5 mm long when dry; style 6-10 mm long, sparsely pubescent; ovary 2-3-locular; ovules 4-8 per locule; fruit globose to pyriform, 7-10 mm long, yellow, the walls thin, slightly spongy, not noticeably glandular;seeds 2-8 in fruits seen, oblong, 3-5 mm long. Type. Dombey s.n., "Rapportepar Dombey. Herb. de Jussieu." (holotype, P, ex herb. Jussieu, n.v.; University of Michigan neg. 1966 of apparentholotype, photo at MICH). Specimens examined. CHILE. Valparaiso:Valle de Marga-Marga (ca. 33?10'S),Sep 1931,Jaffuel

Myrceugenia

& Pirion 3170 (CONC, GH); Las Docas, 7 Mar 1952, Boelcke 6529 (CTES, F, MO), 6538 (F); Vina del Mar, 4 Feb 1923, Behn s.n. (CONC, M, MO); Vina del Mar, Quebrada del Tranque, 18 Aug 1940, Kausel 780 (F, H, HBR, SGO), 782 (F, H, SGO), 783 (H, LP, SGO); Quilpu6, 22 Jan 1931, Behn s.n. (MO, SI, UC); Colliguay, 30 Sep 1973, Zollner 7162 (CTES); Renaca, between Valparaiso and Conc6n, 6 Jan 1952, Poulsen s.n., Poulsen 190 (C); Algarrobo, Quebrada Jer6nimo, 12 Feb 1959, Kausel 4552 (H), 5 Mar 1955, Kausel 4182 (F, H); Algarrobo, Quebrada del Hotel Pacifico, 4 Feb 1957, Kausel 4327 (F); Algarrobo, Isla Negra, Estero el Rosario, 3 Mar 1968, Kausel 5384, 5385 (H); Valparaiso, Jun 1885, Rusby 584 (GH, MICH, MO, PHIL, US), 582 (MICH, PHIL, US); Valparaiso, 1845-1847, U.S. Exploring Expedition-Wilkes s.n. (GH, US); Valparaiso, Didrichsen 3339, 3296 (both C); Valparaiso, Jan 1830, Bertero 1164 (F, GH), May-Jun, Poeppig 144 (F); Valparaiso, Gay s.n. (RB). Santiago: Alhu6, cordillera de la costa, 8 Dec 1938, Kausel 488 (CONC, F, H, SGO); San Antonio, Dec 1926, Claude Joseph 2913 (US). Colchagua: Dep. Santa Cruz, Estero de los Mayus, 100 m, 17 Feb 1946, Kausel 1834 (H, HBR, SGO), 1836 (CONC, H, SGO). Curico: Potrero Grande, Fundo El Pangal, Rio de las Islas, 650 m, 31 Oct 1954, Kausel 4055 (F, H), Oct-Nov 1955, Maige s.n. (F). Talca: Camino longitudinal, puente Rio Claro al norte de Talca, 220 m, 2 Mar 1953, Kausel 3628 (H); Fundo Quebrada del Agua, 700 m, Aug 1936, Grandjot 1503 (MO, SGO); Gualleco, 27 Mar 1971, Landrum & Donoso s.n. (EIF). Maule: Constituci6n, Rio Maule, 28 Jan 1934, Montero 1288 (GH); Constituci6n, Quebrada Honda, 19 Mar 1940, Kausel 721 (F, SGO), 7 Feb 1949, Kausel 2728, 2729 (CONC, F, H, SGO); Constituci6n, Quebrada Camarones, 4 Feb 1949, Kausel 2703 (F, SGO); Camino Cauquenes-Pelluhue, 23 Mar 1971, Landrum & Donoso s.n. (EIF). Linares: Laguna Amargo, ca. 50 km east of Parral, ca. 720 m, 21 Feb 1972, Landrum 925 (MICH); Panimavida, Estero Rari, 17 Mar 1940, Kausel 699 (CONC, F, LP, SGO). Concepci6n: Rio Bio-Bio, Lonco, 23 Mar 1957, Junge 3015 (US); mouth of Bio-Bio river ca. 15 km west of Concepci6n (ca. 36?50'S, 73010'W), 29 Jan 1978, Landrum 3112 (MICH, VALD). Bio-Bio: Salto del Laja, 10 Feb 1953, Behn 13658 (CONC). Malleco: Collipulli, frente a la Genetica, 13 Feb 1947, Ricardi 7474 (CONC).

A lectotype for Eugenia stenophylla [var.] 3 latifolia must be chosen from specimens seen by Berg. Unfortunately I have not seen any of these yet. This species could easily be confused with Myrceugenia pinifolia so they are compared below. M. lanceolata

M. pinifolia

Leaves 2-5 cm long, widest above the middle, oblanceolate to linear Calyx-lobes 1.8-4 mm long, 1-1.7 times as long as wide Hypanthium 2-5 mm long Peduncles solitary or in pairs, uniflorous or bearing 3 flowered dichasia

Leaves 1-3 cm long, widest at about the middie, narrowly elliptic to linear Calyx-lobes 1-2.3 mm long, 0.8-1.3 times as long as wide Hypanthium 1.5-2.5 mm long Peduncles uniflorous and solitary

Myrceugenia lanceolata is also compared to M. hatschbachii in the discussion of that species although there is little chance for confusion because of the geographic separation. The hybridization that frequently occurs between Myrceugenia lanceolata and M. exsucca is discussed in the section on hybridization, and these two species are compared in Key B. In many ways Myrceugenia lanceolata combines characters of M. exsucca and M. pinifolia so one might suspect it of being a species of hybrid origin. But M. lanceolata also has some characteristics found in neither of these species. I think that it is more likely then that M. pinifolia and M. lanceolata have become similar through parallel evolution due to their adaptation to about the same habitat, i.e. the edges of streams in open areas. These two species, M. glaucescens in the southern part of its range, and M. hatschbachii all have strongly discolorous, narrow leaves, often have a reddish pigment in the twigs and all grow in similar habitats. Perhaps this combination of characteristics is adaptive. The darkly colored leaves could be due to the same reddish pigment found in the twigs, which may reflect the warm end of the light spectrum. This characteristic and the

Flora Neotropica

narrowleaves would aid these species in surviving dry periods or exceptionally sunny days. Myrceugenialanceolata grows in Chile from Valparaisoto Malleco. A map of its distributionis shown in Fig. 9. It flowers mainly in Februaryand Marchand the fruits probablymaturemostly in Septemberand October. 7. Myrceugenia rufa (Colla) Skottsberg ex Kausel, Lilloa 13: 134. 1948 ("1947"). Figs. 9, 28A. Myrtusrufa Colla, Mem. Acad. Sci. Torino37: 66 (preprintsMay 1833)late 1834. Eugeniaferruginea Hooker et Arnott, Bot. Misc. 3: 319. 1 Aug 1833.Type. Bridges323 "Valparaiso," 1832(hereby designatedas lectotype, E-GL; Field Museumneg. 31570of isolectotype at W, photo at MICH);Cuming719 (paratype,E-GL). Eugenia rufa (Colla) Barneoudin Gay, Fl. chil. 2: 388. 1847. Lumaferruginea (Hooker et Arnott)A. Gray, U.S. Expl. Exp. 15: 542. 1854. Myrceugeniaferruginea (Hooker et Arnott)Reiche, Anales Univ. Chile 98: 710. 1897;Flora de Chile 2: 294. 1898. Luma rufa (Colla) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 524. 1941.

Shrub 1-2 m high; hairs reddish-brownto whitish, symmetricallydibrachiate, appressed; twigs densely pubescent when young, the hairs persisting until the first bark falls, the originalbark splittingin a reticulatefashion, whitish-grey,the new inner bark at first reddish-brown,becoming whitish-grey; leaves densely pubescent beneath, puberulentabove, sometimes glabrescent with age, broadly to narrowlyelliptic, ovate or oblong, 0.5-1.8 cm long, 0.2-0.5 cm wide, 1.2-4.5 times as long as wide, the marginrevolute;petiole densely pubescent, sometimes glabrescentwith age, 1-2 mm long, not noticeably channeled; veins all indistinct above and below, or the midvein faintly visible; blades light bluish-green to yellowish-green, often lustrous above, reddish-brownto whitish-yellow below, quite coriaceous; peduncles uniflorous,slightly flattened, about 0.5 mm wide, 14 mm long, densely pubescent, solitary or 2-3 in a row in the axils of leaves; bracteoles ovate to broadly oblong, 0.9-1.5 mm long, 0.7-1 mm wide, 1-2 times as long as wide, coriaceous, densely pubescent without, sparsely pubescent within, claspingthe hypanthium,calyx-lobes ovate to suborbicular,stronglyconcave, 1.3-2.6 mm long and wide, usually 0.8-1 times as long as wide, densely to sparsely pubescent without, sparsely pubescent to glabrouswithin; hypanthiumobconic, densely pubescent, ca. 2 mm long; disk 2-2.5 mm across, densely pubescent; style 5-6 mm long, sparsely pubescent; stamens 60-100, 3-6 mm long; anthers 0.4-0.6 mm long when dry; petals more or less orbicular,2-3 mm in diam., very sparselypubescent withinand without;ovary 2-4-locular; ovules 5-13 per locule; fruit 4-8 mm in diam., purplish-black,pubescent; seeds unknown. Type. Bertero s.n. (apparent holotype Bertero 176c or 1765 at TO, n.v., photo

of this specimen at TO, seen; possible isotypes Bertero 1765 F, GH).

Specimensexamined.CHILE. Prov. Coquimbo:Ovalle, CerroTalinaySur (30?50'S,71?37'W),700 m, 23 Oct 1966, Jiles 5017 (CONC); QuebradaAmolanas, en la costa, 125 km al surponientede Ovalle, 1 Nov 1944, Jiles s.n. (H); Pichidangui,Cerro Silla de Gobernador,Sep 1953,Bernaths.n. (F). Aconcagua:Zapallar,Canelilla, 12 Oct 1948, Kausel 2612 (F, H), Kausel 2616 (F); Zapallar, PuntaPite, 80 m, 12 Oct 1948, Kausel 2607 (F); Zapallar,CerroLa Cruz, 10 Oct 1948, Kausel 2584 (F); Zapallar,CaminoPalos Quemados,Aguas Claras, 120 m, 9 Oct 1948, Kausel 2571 (CONC, H, SGO);Papudo,40 m, 19 Sep 1937, Montero3192 (GH). Valparaiso:Valparaiso,1829,Bertero 1170 (F), 1876, Dessauer s.n. (M); Valparaiso,withoutdate, Chamissos.n. (F), Mertenss.n. (GH), 1834, Gaudichaud 234 (F), 1838-1842, U.S. South Pacific Exploring Expedition s.n. (GH, NY, US); Quil-

pu6, en el cerro Chivato, HaciendaPalmas, 12 Sep 1973,Zollner 7129 (CTES);Curauma,Sep 1926, Kausel 273 (H); Curaumilla,Apr 1930, Anwandters.n. (SI); LagunaVerde, 3 Oct 1972, Landrum

944 (MICH); Mirasol, ca. 36 km north of San Antonio (ca. 33?20'S, 71?40'W), 20-30 m, 26 Mar 1978,

Myrceugenia S

-1*. rufesoens . .

0.st

f" S

FIG. 9. Distributions of Myrceugenia rufescens, M. campestris, M. exsucca, M. lanceolata, M. fernandeziana, M. schultzei, and M. rufa.

Landrum 3424, 3425, 3426, 3427 (MICH, SGO); Mirasol, 21 Mar 1943, Bernath 1248 (F), 14-26 Feb 1950, Kausel 3090, 4 Mar 1952, Boelcke 6525 (CTES, MO); Algarrobo, 8 Aug 1959, Kausel 4658 (H),

Sep 1961, Kausel 4624 (H), 20 Sep 1963, Kausel 4673 (H); Algarrobo, La Puntilla, 17 Aug 1958, Kausel 4414 (H), 1 May 1966, Kausel 5019 (H); Algarrobo,Quebradade Guaiquen, 3 Mar 1968, Kausel 5381a (H); Algarrobo,lomajes cerca de la canchade aviaci6n, 6 Mar 1955, Kausel 4185 (H); Conc6n, withoutdate, Poeppig 143 (F). Santiago:El Tabo, 28 Dec 1971,Landrum945 (EIF, MICH); Cartagena, Jan 1920, Claude Joseph 963 (US).

Poeppig 114 (F) supposedly from Talcahuano (Concepcion) and Germain s.n. (F) from "cordillera Maule" are probably wrongly labelled. No modem collections have been made that far south. Myrceugenia rufa is known only from the coast of central Chile, a region that receives little or no rain for most of the year but which does have the benefit of almost constant humid breezes from the ocean. This species usually does not

Flora Neotropica

reach more than a few hundredmeters inland, apparentlyneeding these oceanic winds. A map of its distributionis shown in Fig. 9. The phylogenetic position of Myrceugeniarufa is not clear. It is superficially similar to M. euosma because its leaves are small and densely covered with dibrachiatehairs beneath, but these may be parallel adaptationsto xeric environments. Myrceugeniarufa is easily distinguishedfrom all other species by its smallthick leaves that show little or no venation and that have revolute margins.It flowers fromAugustto October.Its fruitsare so efficientlyattackedby an insect (perhapsa curcurlionid)that I have never been able to find any seeds in them. Whennormal fruits would matureI do not know, but those attacked by insects may persist on the plants at least until March. 8. Myrceugeniaovata (Hooker et Arnott) Berg, Linnaea 30: 670. 1861. Eugenia ovata Hooker et Arnott, Bot. Misc. 3: 319. 1833.

Shrub or small tree up to ca. 8 m high; hairs whitish to reddish-brown,a mixture of simple and dibrachiate;twigs glabrous to densely pubescent when young, glabrescentwith age, the young bark smooth, grey or light brown; leaves glabrousto sparselypubescent beneath, entirelyglabrousto puberulentalong the midvein above, narrowlyto broadly elliptic, ovate or obovate, 0.5-7.5 cm long, 0.2-3 cm wide, 1.2-3.5(-4) times as long as wide; apex acuminate,acute, obtuse or less often rounded; base acuminate,acute, cuneate or truncate;petiole channeled, densely to sparsely pubescent, 1-5 mm long, 0.5-1 mm thick; midvein impressed for entire length or only proximallyabove, moderatelyprominentbelow; lateral veins indistinguishableor up to ca. 10 pairs faintly visible; marginal veins equallinglateralsin prominence;blades grey-green,reddish-brown,or yellow-greenabove, lighterbelow, coriaceous to submembranous,the upper surface dull or lustrous; peduncles uniflorous, flattened, 0.2-2.5 cm long, 0.2-0.5 mm wide, densely pubescent to glabrous, solitary or in pairs in the axils of leaves or bracts; bracteoles ovate to lanceolate, 0.5-1.8 mm long, 0.3-1 mm wide, 1.2-3.8 times as long as wide, sparsely pubescent to glabrous within and without, subcoriaceous to membranous,clasping the hypanthium;calyx-lobes triangularto ovate, 0.8-2.5(-3) mm long and wide, 0.7-1.4 times as long as wide, sparsely pubescent to glabrous within and without, subcoriaceous to membranous,concave; hypanthium densely pubescent, obconic, 0.8-2(-2.5) mm long, the sides straightor slightly convex; disk ca. 1-2.5 mm across, sparsely pubescent; style 4-8 mm long, sparsely pubescent to glabrous; stamens 40-125, 3-8 mm long; anthers 0.2-0.4 mm long when dry; petals glabrous, suborbicular,1.5-4 mm in diam.; ovary 2-4-locular; ovules 2-11 per locule; fruit globose, ca. 4-7 mm in diam., orange to purple-black,few seeded, the seeds oblong, ca. 3-5 mm long. Myrceugenia ovata as circumscribedhere is an inclusive species with both eastern and western populations.In general, specimens from eastern and western South America are distinguishablefrom each other but perhaps 10%of the time one cannot be sure on morphologicalgrounds,from which side of South America a specimen has come. Therefore both populations have been united under one name. In western South America there are two fairly well markedvarieties, one restricted to the Chilean coastal cordillera and coast (Myrceugenia ovata var. ovata) and the other found mainly in the Andes (M. ovata var. nannophylla). Both are found on the island of Chiloe wherethey possibly act as separatespecies, but towards the north in Linares and Nuble, intermediateplants are found that

Myrceugenia

Kausel interpreted as a third species, M. valientei. These northern populations are also those which are most similar to one of the Brazilian varieties, M. ovata var. gracilis. Perhaps as M. ovata migrated southward in western South America after the Pleistocene glaciation it developed into coastal and Andean races which are now meeting in Chiloe. Further investigation of the breeding relationships of those two Chilean varieties would be interesting. In eastern South America Myrceugenia ovata has developed two varieties, one of which, M. ovata var. acutata may be of hybrid origin. This possibility is considered in more detail in the discussion of M. glaucescens. Key to Varieties of Myrceugenia ovata 1. Pedunclesoften in pairs, often subtendedby bracts;leaves mainlyelliptic, 2.7-5 cm long; d. var. acutata. twigs moderatelypubescentto glabrous;Brazil. 1. Peduncles solitary, not subtendedby bracts;leaves 0.3-4 cm long, ovate to elliptic; twigs usually densely pubescent;Brazilto western South America. 2. Calyx-lobesaveragingless than 1.5 mm long; leaves usually averagingover 1 cm long, usually elliptic; hypanthiumgenerallyunder 1 mm long; ovules usually averagingfewer than 6 per locule; Brazil. c. var. gracilis. 2. Calyx-lobes averagingover 1.5 mm long or the leaves averagingless than 1 cm long; leaves usuallyovate; hypanthiumusuallymore than 1 mmlong; ovules usuallyaveraging more than 6 per locule; western South America. 3. Leaves ovate to elliptic, mainlyunder 1 cm long, rarely surpassing1.5 cm long, the b. var. nannophvlla. apex obtuse to rounded;mainlyAndean. 3. Leaves ovate, mainly 1-2 cm long, often surpassing2 cm, the apex acute to acumina. var. ovata. ate; along the coast and in the coastal mountainsof Chile.

8a. Myrceugenia ovata var. ovata.

Figs. 10, 28B.

Eugenia ovata Hooker et Arnott, Bot. Misc. 3: 319, as to type, 1833.

Eugenia cumingii Hooker et Arnott, Bot. Misc. 3: 319. 1833. Type. Cuming31 "Chiloe" (holotype, E-GL; isotype, K). Eugenia trichocarpa Philippi, Linnaea 28: 688. 1858. Type. apparentlyKrause s.n., "Prope Corral" (probable holotype, SGO-052797; probable isotype, G).

Luma ovata (Hooker et Arnott)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 524. 1941. Luma cumingii(Hooker et Arnott)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 524. 1941.

Twigs densely pubescent; leaves mainly ovate, 0.6-4 cm long, 0.5-2 cm wide, ca. 1.2-2.2 times as long as wide; apex acute to acuminate; peduncles ca. 3-15 mm long, 0.3-0.5 mm wide, densely pubescent, solitary in the axils of leaves; calyx-lobes 1.1-2.5(-3) mm long, 1.4-2.5(-3) mm wide, 0.8-1.3 times as long as wide; hypanthium 1-2 mm long; stamens 50-130; ovules 5-11 per locule. Type. Cuming 30, "Chiloe" (holotype, K). Specimensexamined.CHILE. Arauco:Isla Mocha, QuebradaLos Chillanejes,Oct 1938,Bernath s.n. (CONC, H). Valdivia:without specific locality, 1862, Philippi s.n. (G, MICH);Valdivae montibus, Philippi69 (US); Corral,Hohenacker555 (MICH);hills north of Corral,24 Dec 1935, West

4884 (GH, MO, UC); Niebla, ca. 14 km west of Valdivia (ca. 39?45'S, 73?30'W), near sea level, 15 Feb 1978, Landrum 3205, 3206, 3213 (MICH, VALD); Corral, Quitaluto, 450 m, 19 Jan 1931, Gunckel 1941, 1959 (GH, H); Pucatrillo, westlich S. Juan an der Kiiste, 11 Jan 1948, Sparre 3953 (H); Llancacura, 15 Apr 1969, Mahu 2014 (H); Bima district, west of La Uni6n, Cordillera de Alerce, 500-900 m, 11 Feb 1958, Eyerdam 10656 (F, GH, MICH, MO, NY, UC); Amargos, 20 m, 14 Jan 1934, Montero 1337 (GH). Llanquihue: Maullin, Huantrunes, Jan 1941, Pfister 4821 (H). Chilo6: without specific locality, King s.n. (K); Ancud, near Airport, ca. 100 m, 7 May 1972, Landrum 939, 940 (EIF,

MICH); 8 km south of Ancud, 7 May 1972, Landrum876 (MICH);Chepu, 50 m, 10 May 1972, Landrum900 (MICH); Quemchi (ca. 42?10'S, 73?30'W),ca. 50 m, 10 Feb 1978, Landrum3181,

3184, 3194, 3195 (MICH, VALD). Without specific locality: 1833, Bridges s.n. (F), 1870, Schazinann s.n. (G).

This variety is normally an understory tree in the forests along the coast and

Flora Neotropica

in the coastal mountains of Chile. A map of its distribution is shown in Fig. 10. In pastured land it becomes a dense shrub. It flowers from December to February and while the fruits are still unknown it seems probable that they mature from July to November. 8b. Myrceugenia ovata var. nannophylla (Burret) Landrum, Brittonia 32(3): 374. 1980. Fig. 10. Luma nannophyllaBurret,Repert. Spec. Nov. Regni Veg. 50: 50. 1941.

Myrceugenia montana Kausel, Revista Argent. Agron. 9: 55. 1942. Type. Looser 2733 "Termas

de Tolhuaca"(holotype, H).

Mvrceugenia valientei Kausel, Revista Argent. Agron. 11: 323. 1944. Type. Kausel 1336, Chile,

Prov. Nuble, San Fabian"LagunaEl Valiente" (holotype, H). Mvrceugenianannophylla(Burret)Kausel, Revista Argent. Agron. 18: 233. 1951.

Twigs densely pubescent; leaves mainly ovate, elliptic towards the northern edge of range, 3-16 mm long, 2-8 mm wide, 1.2-2.6 times as long as wide; apex obtuse to rounded; peduncles ca. 2-15 mm long, 0.3-0.5 mm wide, solitary in the axils of leaves; calyx-lobes 1-2.2 mm long, 1-2 mm wide, 0.8-1.4 times as long as wide; hypanthium 1-2 mm long; stamens 50-110; ovules 3-8 per locule. Type. Philippi s.n., "Chile: Prov. Valdivia, Quellgebiet des Rio Futa, in den Alerzale" (holotype, B, apparently lost). Specimensexamined.CHILE. Linares:LagunaAmargo,Bullileo, ca. 50 km east of Parral,ca. 720 m, 21 Feb 1972, Landrum871 (MICH). Nuble: San Carlos, Feb 1926, Claude Joseph 3971 (US); Caminoa la LagunaEl Valiente, 800-900 m, 6 Mar 1944,Kausel 1333(H); LagunaEl Valiente, 10001100m, 6 Mar 1944,Kausel 1335, 1337(bothat H). Malleco:Cordillerade Nahuelbuta,Coyancahuin, ca. 500 m, 20 Feb 1939, Kausel 579 (H, CONC),600 m, 14-16 Feb 1939, Kausel541 (CONC,H, LP, SGO), 19-20 Feb 1939, Kausel 581 (F), 700 m, 14-16 Feb 1939, Kausel 542 (F), 1000m, 14-16 Feb 1939, Kausel 549 (F, SGO); Bafos de Tolhuaca, 1000 m, 28 Feb 1947, Kausel 2442 (F, H, HBR, SGO);LagunaVerde, Tolhuaca, 1250m, 28 Feb 1947,Kausel2817 (F, SGO), Kausel 2818 (F); RaroRuca, a orillas del rio Cautin, 15-25 Feb 1947, Kausel 2370 (CONC, F, H, SGO); Curacautin,rio Mantible,490 m, 17 Feb 1947,Kausel2373 (F, SGO);TermasManzanar,ca. 25 km east of Curacautin, 10 Jan 1973, Landrum937 (EIF, MICH).Cautin:Temuco, 19 May 1927, ClaudeJoseph 4880 (US); Lago Villarrica,BaniosPalguin,ca. 800 m, 9 Feb 1965,Kausel4978, 4882 (both H). Valdivia:without specific locality, 1862, Philippi s.n. (MICH). Osorno: Nadi entre Osorno y Puyehue, 19 Feb 1942, Kausel 1049 (H, SGO);bog along road to Antillanca,ParqueNacional Puyehue, 25 Feb 1971, Landrumn935 (EIF, MICH). Chilo6:Tirunquina[?], 19 Feb 1932, Junge 309 (CONC, H), 6 Feb 1932, Gunckel3492 (H); at corner of main Ancud-Castroroad and road to Chepu, 8 May 1972, Landrum 929 (MICH);14 km northof Castroalong highwayto Ancud, ca. 200 m, 9 Feb 1978, Landrum3169, 3172, 3175, 3176 (MICH, VALD).

ARGENTINA.Neuquen:Dpto. Huiliches, Lago de Epulafqu6n,5 Feb 1963, Movia s.n. (CTES); Lago Quillen, 4 Feb 1968, Eskuche& Klein 0100 (CTES, SI); Hua-Hum, 14 Dec 1937, Kalela 1450 (H); camino Trafula Ruca Malen, 26 Feb 1953,Boelcke & Correa7248 (MICH);Lago Correntoso, Ruca Malen, Feb 1944, Bernasconis.n. (SI); Lago Correntoso,Quintapuray,800 m, 26 Mar 1934, Spegazzini s.n. (MICH); Lago Correntoso, 6 Nov 1937, Kalela 687, 713 (H), 20 Mar 1939, Cabrera

5031 (GH, LP); Lago Espejo, 20 Mar 1939,Cabrera5038 (GH, LP), 1 Mar 1978,Landrum3290, 3291 (LP, MICH);Lago Espejo, rio Campana,8 Feb 1940, Cabrera6006 (H, NY); Villa La Angostura, 5 Feb 1959, Boelcke 10551 (CTES), Feb 1940, Perez Moreau s.n. (H); Villa La Angostura (ca. 40?50'S,

71?50'W),ca. 800 m, 27 Feb 1978, Landrum3283, 3284, 3285 (LP, MICH);Cumelennear Villa La Angostura, 18 Feb 1952, Pedersen 1544 (C); Puerto Manzanonear Villa La Angostura,ca. 800 m, 1 Mar 1978, Landrum3292, 3294 (LP, MICH);PeninsulaQuetrihu6,2 May 1942, Diem 359 (H).

Although no type has been found for Luma nannophylla, it is very likely that it belongs to this variety judging from the protolog. Myrceugenia ovat6 var. nannophylla, mainly an Andean entity, typically inhabits cool areas along lakes and streams and in bogs. A map of its distribution is shown in Fig. 10. It often grows with Nothofagus antarctica in what appear to be cold air pockets. In the region of Lago Nahuel Huapi, it hybridizes with M. exsucca, a subject which will be discussed in the section on hybridization.

Myrceugenia

Myrceugenia ovata var. nannophylla flowers from February to April and the

fruits seem to matureabout 12 months later. It is comparedwith M. pinifolia in the discussion of that closely related species. 8c. Myrceugenia ovata var. gracilis (Burret) Landrum, Brittonia 32(3): 374. 1980. Fig. 10. Luma gracilis Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 534. Eugenia regnelliana Berg, in Mart. Fl. Bras. 14(1): 245. 1857. Type. Regnell II n. 120 (hereby

designatedas lectotype; lectotype, MEL, n.v.; isolectotype, SP; Field Museumneg. 20973 of isolectotype at C, photo at MICH);Widgren556 (paratype,MEL, n.v.; isoparatype,US) "Habitatin prov. Minarumad Caldas,""(v. in hb. Sonder.sub nomineE. obtusifoliaCamb. Miq.)." Eugenia itatiaiensis Kiaerskou,Enum. Myrt. bras.: 166. 1893.Type. Glaziou6545 "In summo monte Itatiaiaprope Ribeiraom. Januarii1873florentem"(holotype, C; Field Museumneg. 20953of holotype, photo MICH). Luma regnelliana(Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 535. 1941. Luma itatiaiensis (Kiaerskou)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 534. 1941. Luma macromischaBurret, Repert. Spec. Nov. Regni Veg. 50: 53. 1941.Type. Loefgren s.n., "S. Paulo, S. Francisco," "Mus. S. Paulo n. 3552" (holotype, SP). Luma oreophila Burret, Repert. Spec. Nov. Regni Veg. 50: 54. 1941. Type. Lutzelburg6550, "Staat Rio de Janeiro,Serrados Orgaos, 2400 m, MorroAssu" (isotype, M). Myrceugeniaregnelliana (Berg) Legrandet Kausel, in Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 11. 1953. Mvrceugeniaitatiaiensis (Kiaerskou)Legrandet Kausel, in Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 12. 1953. Mvrceugenia regnelliana var. leptophylla Legrand, Darwiniana 11(2): 354. 1957. Type. Reitz

2576, SantaCatarina,Campodos Padres, 1900-2000m (holotype, MVM;isotypes, H, HBR, PACA). Myrceugeniaregnelliana var. gracilis (Burret)Legrand,Darwiniana11(2):354. 1957. Myrceugenia regnelliana var. capillaris Legrand, Sellowia 13: 303. 1961. Type. Reitz & Klein

7199, Santa Catarina,Lauro Muller-Urussanga:Pinheiralda Companhia,Itanema, 300 m, 20 Sep 1958(holotype, MVM;isotypes, H, HBR).

Mvrceugenia regnelliana var. coriacea Legrand, Sellowia 13: 303. 1961. Type. Reitz 2413, Santa

Catarina,Bom Retiro, Campodos Padres(holotype, MVM;isotypes, H, HBR).

Mvrceugenia regnelliana forma itatiaiensis (Kiaerskou) Legrand, in Legrand et Klein, Flora

Ilustr. Catar. [Mirt]:426. 1970.

Twigs densely to sparsely pubescent; leaves 1-3 cm long, 0.5-1 cm wide, 1.73 times as long as wide; apex acute, acuminateor obtuse; peduncles 2-15 mm long, 0.2-0.3(-0.4) mm wide, glabrousto densely pubescent, solitary in the axils of leaves; calyx-lobes (0.6-)0.8-1.5 mm long and wide, 0.7-1.4 times as long as wide; hypanthium0.8-1.8 mm long; stamens 40-110; ovules 2-8(-11) per locule. Type: "Agulhas Negras, S. Paulo (Mus. Paulista n. 435, Secc. Bot. n. 26994 com. F. C. Hoehne)" (holotype, SP). Specimens examined. BRAZIL. MinasGerais:Aiuruoca,Pico do Papagaio,23 Jan 1886, Glaziou 16073 (P, C, G); Aiuruoca, Nov 1897, Silveriras.n. (R); Delfim Moreira, a 7 km de Is6polis, na estrada para S. Francisco, 14 Oct 1968, J. Mattos 15385 (SP). Rio de Janeiro:ParqueNacional da Serrados Orgaos, Campodas Antas, 2000 m, 3-5 Sep 1949, Brade 20053 (MVM, RB); Terez6polis, Posse, morrodas antenasde televisao, 10 Feb 1968, Sucre 2325 (MBM, RB); Itatiaia,planalto,2000 m, Jan 1938, Barreto & Brade 16033 (MBM, MVM, RB), 2550 m, 5 Feb 1969, Sucre 4677 (RB, US);

Itatiaia, Estacao Biol6gica, 22?25'S,44?41'W,2200 m, 7 Jan 1929, L. B. Smith 1734 (US); Itatiaia, km 15-16, 15 Feb 1935, CamposPorto 2787 (MICH);Itatiaia,Beira do rio CampoBello, 900 m, 27 Mar 1942, Brade 17295 (RB); Rio das Flores, 31 Jan 1935, Campos Porto 2716 (MICH, MVM, RB);

Agulhas Negras, 2800 m, 27 May 1935, Brade 14608 (LP, MICH, MVM, RB). Sio Paulo: Campos

do Jordao, Fazenda da Guarda, 1600 m, 25 Nov 1949, Kuhlmann 2185 (SP); Campos do Jordao,

Fazendada Guarda,26 Aug 1967,J. Mattos 15051(SP); Camposdo Jordao,Reserva Florestal, 1600

m, 5 Dec 1977, Landrum 2826, 2827, 2828 (MICH, SP), 2000 m, Landrum 2815 (MICH, SP); Campos

de Jordao,ParqueEstatal, 23 Apr 1977,J. Mattos 15798(SP); Paranapiacaba,Estacao Biol6gica, 27 Dec 1966, J. Mattos 14376 (SP); Itarare,Camposde Sao Pedro, Fazenda Ventania, Boca da Serra

Flora Neotropica

de Bom Sucesso, ca. 1000 m, 10 Oct 1966, J. Mattos 14900 (SP). Parana: Mun. Bocaiuva do Sul, Cabeca d'Anta, 21 Dec 1960, Hatschbach 7595 (MBM, MVM); Mun. Campina Grande do Sul, Serra Capivari Grande, 15 Jan 1969, Hatschbach 20745 (MICH), 1500 m, 8 Feb 1971, Hatschbach 23405 (C, CTES, HB, MBM, MICH, MVM, UC); Mun. Campina Grande do Sul, Pico Caratuva, 1950 m, 8 Feb 1968, Hatschbach 18568 (MICH, MVM), 2 Aug 1967, Hatschbach 16823 (MICH); Mun. Campo Largo, arredores, 950 m, 3 Oct 1958, Hatschbach 5050 (MBM, MVM); Mun. Castro, Fundao, 2 Oct 1964, Hatschbach 11646 (MBM, MVM); Mun. Colombo, Pogo Preto, 10 Jan 1963, Hatschbach 9693 (MBM, MVM); Mun. Guarapuava, Serra da Esperanca, 20 Oct 1960, Hatschbach 7336 (HB, HBR, MBM, MICH, MVM); Mun. Guaratuba, Serra de Aragatuba, 1350 m, 31 Jan 1960, Hatschbach 6692 (MBM, MVM); Ipiranga, Monte Alegre, 8 Feb 1904, Dusen 3377 (R, US); Mun. Lapa, road to Balsa Nova, 7 Nov 1977, Landrum 2455 (MBM, MICH); Mun. Morretes, Pico Olimpico, 1547 m, 15 Jan 1950, Hatschbach 1744 (MBM, MICH, MVM); Mun. Pirai do Sul, Serra das Furnas, 30 Mar 1957, Hatschbach 4021 (PACA, MBM, MICH, MVM); Mun. Rio Negro, Pangare, 29 Nov 1956, Hatschbach 3412 (HBR, MBM, MICH, MVM); Mun. Sao Joao do Triunfo, Sao Joao do Triunfo, 7 Feb 1967, Hatschbach 17708 (C, MBM, MICH, MVM, UC); Mun. Tijucas do Sul, Saltinho, 5 Nov. 1956, Hatschbach 3403 (MBM, MVM), 25 Oct 1971, Hatschbach 27584 (C, HB, MBM, UC). Santa Catarina: Mun. Bom Jardim, Curral Falso, 1500 m, 10 Dec 1958, Reitz & Klein 7774 (H, HBR); Mun. Bom Jardim, Fazenda da Laranja, 1400 m, 13 Dec 1958, Reitz & Klein 7870 (HBR, H); Mun. Bom Retiro, Campo dos Padres, 2000 m, 20 Dec 1948, Reitz 2574 (H, HBR), 1900 m, 17 Dec 1948, Reitz 2425 (H, HBR); Mun. Bom Retiro, 930 m, 9 Jan 1948, Reitz 1985 (H, MICH, UC); Mun. Bom Retiro, Riosinho, 1000 m, 23 Dec 1948, Reitz 2741 (H, HBR, PACA, R), 24 Dec 1948, Reitz 2769 (H, HBR); Mun. Bom Retiro, forest between Fazenda Santo Antonio and Fazenda Campo dos Padres, 14001650 m, 21 Nov 1956, Smith & Klein 7793 (HBR, R, US); Mun. Bom Retiro, forest between Fazenda Santo Antonio and the fall of Rio Canoas, 1300-1400 m, 22 Nov 1956, Smith & Klein 7835 (HBR, R, US), Smith & Klein 7842 (HBR, US); Mun. Bom Retiro, dwarf forest, Fazenda Campo dos Padres, 1650 m, 17-19 Nov 1956, Smith, Reitz & Klein 7731 (HBR, US), 7732 (GH, HBR, R, RB, UC, US), 7733 (HBR, R, US), 22 Jan 1957, Smith, Reitz & Klein 10297 (HBR, R, US); Mun. Bom Retiro, Campina, Riozinho, 1000 m, 24 Nov 1956, Smith & Klein 7922 (C, HBR, R, US); Mun. Cacador, Imbuial, 800 m, 7 Dec 1962, Klein 3448 (H, HBR), 8 km north of Cacador, 950-1100 m, 21 Dec 1956, Smith & Reitz 8955 (HBR, MICH, R, US), Fazenda Carneiros, northeast of Cacador, 950-1100 m, 21 Dec 1956, Smith & Reitz 9014 (HBR, R, US); Mun. Campo Alegre, Morro Iquererim, 1500 m, Smith & Klein 8519, 8533 (HBR, R, US), 1400 m, 9 Jan 1958, Reitz & Klein 6051 (H, HBR, UC, US), 1000-1300 m, 8 Nov 1956, Smith & Klein 7361 (F, HBR, MO, R, RB, US), Smith & Klein 7385 (HBR, R, US), 900 m, 18 Oct 1957, Reitz & Klein 5192, 5197 (HBR, US); Mun. Lajes, Lajes, 850 m, 25 Sep 1960, Cabrera, Reitz & Klein 9964 (H, HBR); Mun. Lajes, 11 km east of Otacilio Costa, ca. 1000 m, 21 Nov 1977, Landrum 2637 (MBM, MICH); Mun. Lauro Miller, Novo Horizonte, 400 m, 19 Sep 1958, Reitz & Klein 7224 (H, HBR); Mun. Lebon Regis, Rio dos Patos, 900 m, 6 Dec 1962, Klein 3388 (H, HBR); Mun. Mafra, Mafra, 5 Dec 1948, Hatschbach 1142 (MBM, MICH); Mun. Matos Costa, Matos Costa, 1000 m, 27 Oct 1962, Reitz & Klein 13735 (H, HBR); Mun. Papanduva, north of the Serra Geral on the Estrada de Rodagem Federal, km 193, 700-900 m, 6-7 Dec 1965, Smith & Klein 8394 (HBR, R, US); Mun. Papanduva, Serra do Espigao, 1000 m, 14 Dec 1962, Klein 4010 (H, HBR, UC); Mun. Papanduva, Lajeadinho, 750 m, 13 Dec 1962, Klein 3968 (H, HBR, MBM, US); Mun. P6rto Uniao, 800 m, 9 Dec 1962, Klein 3635 (H, HBR), Sao Miguel, 800 m, 9 Dec 1962, Klein 3625 (H, HBR, MBM, US); Mun. Sao Francisco do Sul, Morro do Campo Alegre, 1200 m, 21 Dec 1960, Reitz & Klein 10466, 10490 (both H, HBR, UC, US); Mun. Sao Jose, Serra da Boa Vista, 1000 m, 10 Nov 1960, Reitz & Klein 10417 (H, HBR), 27 Dec 1960, Reitz & Klein 10592 (H, HBR, UC), 700 m, 13 Apr 1961, Reitz & Klein 10993 (H, HBR). Rio Grande do Sul: Mun. Cambara, Fortaleza, ca. 1100 m, 27 Dec 1977, Landrum 2984 (MICH, PACA).

The type of Luma gracilis was collected by H. Luederwalt according to the label but Burret makes no mention of this. This entity has gone under the name Myrceugenia regnelliana since 1953 but that epithet does not have priority at varietal rank. Myrceugenia ovata var. gracilis grows in the foggy forests of the eastern planalto and as an understory tree in Araucaria forests. A map of its distribution is shown in Fig. 10. It flowers mainly from October to January and the fruits probably mature about 8 to 12 months later. Both M. hatschbachii and M. smithii appear to be close relatives and are perhaps derived from ancestral stock similar to M. ovata var. gracilis. Both have much narrower leaves and are stream side shrubs of more or less open situations, M. hatschbachii of the campos of the planalto and M. smithii of the lowlands east of the planalto. Myrceugenia hatsch-

Myrceugenia

bachii has very little pubescence on the twigs and peduncles and has peduncles normally longer than in M. ovata var. gracilis. Myrceugenia smithii has peduncles shorter than usual in M. ovata var. gracilis and its twigs often have an orange-brown, flaky bark. 8d. Myrceugenia ovata var. acutata (Legrand) Landrum, Brittonia 32(3): 374. 1980. Fig. 10. Myrceugenia acutata Legrand, Darwiniana 11(2): 351. 1957.

Twigs moderately pubescent to glabrous; leaves elliptic to lanceolate, 2-7.5 cm long, 0.8-3 cm wide, 1.5-4 times as long as wide; apex acuminate to acute; peduncles 3-25 mm long, 0.3-0.5 mm wide, glabrous to sparsely pubescent, solitary or in pairs in the axils of leaves or bracts; calyx-lobes 1-1.7 mm long, 0.9-2 mm wide, 0.6-1.4 times as long as wide; hypanthium ca. 1-1.5 mm long; stamens ca. 50-100; ovules 4-7 per locule. Type. Altamiro & Walter 66, "Brasil: Rio de Janeiro, Itatiaia pr. Campo Belo" (holotype, MVM; isotypes, MICH, MO, RB). Specimens examined. BRAZIL. Parana: Mun. Campina Grande do Sul, Pico Caratuva, 2 Aug 1967, Hatschbach 16857 (MVM); Mun. Campina Grande do Sul, Rio Taquary, 9 Dec 1956, Hatschbach 3579 (MBM, MICH, MVM). Santa Catarina: Mun. Bom Jardim, Curral Falso, 1500 m, 11 Dec 1958, Reitz & Klein 7807 (H, HBR, UC); Mun. Bom Jardim, Serra de Oratorio, 1400 m, 23 Oct 1958, Reitz & Klein 7427 (H, HBR, UC, US); Mun. Bom Jardim, ca. 1400 m, 25 Jan 1971, Eskuche 01470 (MVM); Mun. Governador Celso Ramos, Jordao, 300 m, 18 Oct 1971, Klein & Bresolin 9765 (MVM); Mun. Florian6polis, Morro do Rio Vermelho, 200 m, 11 Sep 1968, Klein & Bresolin 7916 (MVM), 17 Oct 1968, Klein 7936 (MVM); Mun. Palhoca, Morro do Cambirela, 700 m, 20 Oct 1971, Klein & Bresolin 9846 (MVM); Mun. Sao Joaquim, a 10 km de Sao Joaquim, depois da Chapada Seca, 1400 m, 22 Oct 1961, Pabst 6205 (R). Rio Grande do Sul: Mun. Cambara, Tambecinho, ca. 1000 m, 28 Dec 1977, Landrum 3012, 3013, 3014 (MICH, PACA), 3 Nov 1954, Rambo 55958 (PACA), 13 Nov 1953, Ramibo 54483 (PACA); Bom Jesus, Serra da Rocinha, 18 Jan 1950, Rambo 45444 (PACA); Garibaldi, 29 Oct 1957, Camargo 2244 (PACA); Farroupilha, 22 Oct 1956, Carmargo 839 (PACA), 1 Sep 1957, Camnargo 1684 (PACA).

The flowers of this variety are essentially identical to those of Myrceugenia ovata var. gracilis but in the characteristics of twig pubescence, leaf size and the presence of bracts subtending some peduncles it approaches M. glaucescens var. latior. Myrceugenia ovata var. acutata flowers mainly in October and November and apparently the fruits mature shortly thereafter as in M. glaucescens. It is perhaps an entity of hybrid origin as is discussed in more detail under M. glaucescens. Myrceugenia ovata var. acutata is also rather similar to M. cucullata with which it is compared directly in Key L. The distinction between Myrceugenia ovata var. acutata and M. ovata var. gracilis is most difficult in southern Santa Catarina where there are many intermediates. Myrceugenia ovata var. acutata is found from Rio de Janeiro to Rio Grande do Sul. A map of its distribution is shown in Fig. 10. 9. Myrceugenia pinifolia (F. Philippi) Kausel, Rev. Mirt. Chile 2. 1940; Revista Argent. Agron. 9: 54. 1942. Fig. 10. Eugenia pinifolia F. Philippi, Anales Univ. Chile 84: 758. 1893. Mvrceugenia stenophvlla var. pinifolia (F. Philippi) Reiche, Anales Univ. Chile 98: 713. 1897; Flora de Chile 2: 297. 1898.

Shrub up to ca. 2 m high; hairs reddish-brown to whitish, a mixture of simple and dibrachiate; twigs densely pubescent when young, glabrescent with age, the hairs usually spreading; leaves narrowly elliptic to linear, 1-3 cm long, 0.2-0.8

Flora Neotropica

cm wide, 3-12 times as long as wide, glabrous to very sparsely pubescent, the hairs mainlyappressed, dibrachiate;apex and base acute or obtuse; petiole channeled, densely pubescent, 0.5-3 mm long, 0.3-0.8 mm thick; midvein slightly impressed proximally to not at all above, moderately prominent to indistinct below; lateral and marginalveins indistinct;blades coriaceous to subcoriaceous, grey-green,often darkabove, lightgrey-greento light yellow-greenbelow, usually strongly discolorous, the surfaces dull;peduncles uniflorous,5-10 mm long, 0.30.5 mm wide, densely pubescent, solitary in the axils of leaves; bracteoles ovate to lanceolate, (0.5-)0.7-1.8 mm long, 0.3-1 mm wide, 1.3-2.6 times as long as wide, subcoriaceous, glabrous or nearly so within, sparsely pubescent without, clasping the hypanthium;calyx-lobes bluntly ovate to shortly oblong, 1-2.3 mm long, 1-2.5 mm wide, 0.8-1.3 times as long as wide, subcoriaceous, glabrousor nearly so within, sparselypubescentwithout, concave; hypanthiumobconic, 1.52.5 mm long, densely pubescent; disk sparsely pubescent to glabrous, 1.5-2.5 mm across; style glabrous6-7 mm long; stamens 100-140, 3-8 mm long; anthers 0.3-0.4 mm long when dry; petals glabrous, suborbicularca. 3 mm in diam.; ovary 2-locular;ovules 6-12 per locule; fruit unknown. Type. PresumablyPhilippis.n., "Habitatin Araucaria1. d. Curanilahue,florebat mensi decembri" (apparentholotype, SGO-064202). Specimens examined. CHILE. Maule: Constituci6n,QuebradaCamarones,4 Feb 1949, Kausel 2702 (CONC, H); Constituci6n,QuebradaHonda, 7 Feb 1949, Kausel 2730 (CONC, F, H, SGO); Empedrado,31 Jan 1943, Bernaths.n. (H); road from Cauquenesto Pelluhue, Mar 1971, Landrum & Donoso s.n. (EIF). Concepci6n:Villamavida,80 m, 14 Feb 1953, Kausel 3570 (F); Villamavida, 12-25 Feb 1945, Kausel 1590, 1591 (H, SGO), Kausel 1592 (CONC, H, HBR, SGO); about 8 km east

of Concepci6n, ca. 100 m, 27 Jan 1978, Landrum3086, 3090, 3092, 3093 (MICH, VALD), 20 Mar 1978, Landrum 3379, 3381, 3384, 3386 (MICH, SGO); Laraquete, Feb 1937, Kausel 347 (H), 24-25

Mar 1940,Kausel 746 (CONC, F, HBR, LP). Arauco:caminoentre Curanilahuey Cafiete,Quebrada del rio Trongal(37?33'S,73?23'W),170 m, 11 Jan 1972, Quezada, Rodriguez& Weldt2 (CONC); Lago Lleu-Lleu,23-24 Mar 1939, Kausel 602 (SGO). Withoutspecific locality: Dombey 776 (P).

Myrceugeniapinifolia is closely related to and perhaps derived from ancestral stock similarto M. ovata var. nannophylla.The differencesbetween the two are summarizedbelow. M. pinifolia

M. ovata var. nannophylla

Leaves 1-3 cm long, 3-12 times as long as wide Hypanthium1.5-2.5 mm long Ovules per locule 6-12

Leaves 0.3-1.6 cm long, 1.2-2.6 times as long as wide Hypanthium1-2 mm long Ovules per locule 3-8

Myrceugenia ovata var. nannophylla, mainly Andean in distribution, is geo-

graphicallyseparatedfrom M. pinifolia except perhaps in the Cordilleraof Nahuelbuta. Very likely in that region they are separatedaltitudinally,M. pinifolia growing at elevations below 200 m and M. ovata var. nannophylla at about 500 m. The differences between these entities are admittedlysmall but to unite Myrceugenia pinifolia with M. ovata would increase the intraspecific variation of M. ovata and decrease its distinctness from M. parvifolia, M. hatschbachii, M.

smithii and M. leptospermoides. One would have to consider unitingall into one very large, variable species. I believe it preferable to retain M. pinifolia as a separate species in order to minimizeconfusion. Myrceugeniapinifolia is also comparedto two other similarspecies, M. hatschbachii and M. lanceolata, in the discussion of those taxa. Myrceugeniapinifolia, a shrubgrowingalong streams and aroundlakes, flow-

Myrceugenia

ers in January and February. Fruiting specimens are still unknown. A map of its distribution is shown in Fig. 10. 10. Myrceugenia obtusa (A. P. de Candolle) Berg, Linnaea 30: 699. 1861. Figs. 7A, 10, 28C. Eugenia obtusa A. P. de Candolle, Prodr. 3: 266. 1828. Myrtus raran Colla, Mem. Acad. Sci. Torino 37: 66. (preprints May 1833) late 1834. Type. Bertero s.n., "Valparaiso loco dicto las Tablas" (holotype, TO, n.v.; photo of apparent holotype from TO seen). Eugenia raran (Colla) Barneoud, in Gay, Fl. chil. 2: 388. 1847. Luma obtusa (A. P. de Candolle) A. Gray, U.S. Expl. Exped., Phan.: 541. 1854. Myrceugenia chilensis Berg, Linnaea 27: 132. 1856. Illegitimate name, because Eugenia obtusa A. P. de Candolle is cited as a synonym. Eugenia polyantha Philippi, Linnaea 28: 639. 1857. Type. Germain s.n., "In cordillerade Linares dicta in praedio Culmen" (holotype, SGO; Field Museum neg. 31603 of apparent isotype at W, photo MICH). Myrceugenia obtusa [var.] a raran (Colla) Berg, Linnaea 30: 699. 1861. Myrceugenia obtusa [var.] P berteroana Berg, Linnaea 30: 699. 1861. Type. None cited, presumably some Bertero collection (Field Museum neg. 19741 of possible isotype at W, photo at MICH). Myrceugenia obtusa [var.] y polyantha (Philippi) Berg, Linnaea 30: 699. 1861.

Shrub or small tree 2-8 m high; hairs reddish-brown, a mixture of simple and dibrachiate; twigs reddish-brown when young, becoming greyish brown with age, densely pubescent when young, the hairs spreading, persisting until the first bark exfoliates; leaves opposite to subopposite, glabrous or with a few scattered hairs, elliptic to ovate, 1-3 cm long, 0.6-2(-2.5) cm wide, 1-2.2 times as long as wide; apex bluntly acute, obtuse or rounded; base acute, obtuse to rounded (rarely slightly cordate); midvein not noticeably impressed above, moderately prominent to indistinct below; lateral veins indistinct or up to ca. 5 pairs faintly visible; marginal veins indistinct; blades coriaceous, often brittle even when alive, greygreen to brown above, lighter grey-green or brown below, the surfaces dull or lustrous, often wrinkled, with the appearance of shiny paper; peduncles uniflorous, flattened, 2-15 mm long, 0.4-0.6 mm wide, moderately to densely pubescent, solitary in the axils of leaves; bracteoles leaf-like, petiolate, linear, oblong, elliptic or spatulate, 2-6 mm long, 0.5-2.5 mm wide, 2-4.5 times as long as wide, subcoriaceous, glabrous to sparsely pubescent, not clasping the hypanthium; calyx-lobes ovate to oblong, 2-4 mm long, 1.7-2.5 mm wide, 1-1.8 times as long as wide, submembranous, glabrous within, sparsely pubescent to glabrous without, concave; petals suborbicular, 3-4 mm in diam.; hypanthium densely pubescent, 1-2 mm long, more or less obconic; disk 2-2.5 mm across, sparsely pubescent to glabrous; stamens 90-190, 4-8 mm long, anthers 0.2-0.4 mm long when dry; style 5-7 mm long, glabrous to sparsely pubescent; ovary 2-4-locular; ovules 5-9(-12) per locule; fruit dark purple-black, globose to elongate, 5-10 mm in diam.; seeds few, 3-4 in fruits seen, more or less round, 4-5 mm in diam. Type. Dombey s.n., "in Peruvia" (holotype, G-DC, n.v.; Field Museum neg. 7945 of holotype, photo at MICH). Specimens examined. CHILE. Coquimbo: Illapel, Fundo Palo Colorado II, Quebrada del Buitre, 14 Oct 1948, Kausel 2639 (CONC). Aconcagua: Zapallar, quebrada del agua potable, 13 Nov 1938, Kausel 446 (H), 447 (H, LP); Zapallar, Quebrada del Tigre, 27 Feb 1952, Boelcke 6465. Valparaiso: Quintero, 1833, Bertero 1167 (GH, MICH); Caj6n de San Pedro, west slope of Campana de Quillota, 900 m, 19 Jan 1936, West 5192 (MO, UC); faldeos sud Cerro Campana, 13 Jan 1950, Boelcke 3913 (F); Granizo, Caj6n Grande (foot of Cerro Campana), near Olmu6, 15 km east of Limache, 1400 ft, 1 Jan 1966, Meyer 9709 (MICH, MO, UC); Quebrada La Troya at base of Campana, ca. 12 km from Limache, 400 m, 4 Dec 1938, Morrison 16731 (MO, UC); Cerro la Campana, 1100 m, 8 Jan 1939, Kausel 505 (H); Quilpu6, Valle Marga-Marga, Quebrada de los Padres Franceses, 1 Mar 1942, K.

Flora Neotropica

Behn 23109 (CONC);Caj6nde la Pataguilla,ca. 4 km east of Lliu-Lliu,23 Dec 1951,Hutchinson193 (UC, US); Las Docas, 7 Mar 1952, Boelcke 6539 (CTES, MO);Peniuelas,Fundo San Juan, 17 Dec 1940, K. Behn 23110 (CONC);Placilla, HaciendaSan Juan, 4 Feb 1941, Kausel 908 (F); Valparaiso, 1845-1847, Didrichsen 3345 (C); Valparaiso, U.S. Exploring Expedition-Wilkes

s.n. (GH, NY, US);

PlayaMirasol,ca. 36 km northof San Antonio(ca. 33?20'S,71?40'W),20-30 m, 26 Mar1978,Landrum 3421 (MICH, SGO). Santiago:Dep. de San Antonio, Quebradade Venega, 350 m, 2-5 Apr 1942, Pisano & Baraona 1393 (H); San Antonio, Nov 1942, Claude Joseph 2872 (US). O'Higgins: Fundo

Cocalan, Las Cabras, 17 Feb 1947, Sparre2308 (MICH);HaciendaCocalan, QuebradaLas Palmas, 14 Dec 1952, Kausel 3514 (H). Colchagua:Yaquil, ca. 20 km west of San Fernando,ca. 300 m, 21 Apr 1971,Donoso & Landrums.n. (EIF); Dep. SantaCruz, Apalta,HaciendaMillahue,16Feb 1946, Kausel 1808 (H, HBR); San Fernando,ca. 400 m, 12 Oct 1938, Montero3511 (GH); withoutspecific locality, 1888, Philippi 728, 732 (US). Curic6:La Montafia,ca. 50 km northeastof Curic6, 30 Apr 1970,Landrum938 (MICH);Mataquito,Caminode Cutemua Quirahue,150m, 17 Feb 1946, Kausel 1849 (CONC). Talca: cerca Gualleco, 27 Mar 1971, Donoso & Landrums.n. (EIF, SGO). Maule: Coronel, 600 m, 23 Feb 1972, Donoso, Landrum & Gajardo s.n. (EIF). Concepci6n: Tumbes Pe-

ninsula,8 km northof Talcahuano,100m, 4 Jan 1936, West5079 (GH, MO, UC); Yumbel,cercanias de Rere, 6 Jan 1942, Munoz & Johnson 2724 (H); Maule near Coronel, 28 Jul 1917, Skottsberg1478 (F, NY); Dep. Coronel, HaciendaEscuadr6n,25 Mar 1940, Kausel 758 (F, HBR); Coronel, 25 Dec 1968, Merxmuller 24901 (M); Concepci6n, Dec 1930, Jaffuel 1292 (GH), Dec 1931, Jaffuel 2929 (GH);

ParqueHualp6n,ca. 15 km west of Concepci6n(ca. 36?50'S,73?10'W),near sea level, 29 Jan 1978, Landrum 3107, 3113 (MICH, VALD). Arauco: Canete, Lago Lanalhue (37?55'S, 73?20'W), 25 m, 14 Dec 1974, Rodriguez 644 (CONC); Lanalhue, Claude Joseph 5958 (US). Malleco: Mininco, 4 Dec

1952, Kunkel407 (C). Cautin:Cerro Nielol, Temuco, 15 Mar 1978, Landrum3363 (MICH, SGO). Withoutspecific locality: Poeppig 76 (F, GH), Pavon 890 (F).

Myrceugeniaobtusa is a common species of the mediterraneanregionof central Chile but extends as far south as Temuco. A map of its distributionis shown in Fig. 10. It is probably a derivative of M. ovata or a not too distant common ancestor. It is easily distinguishedfrom all the other species in the genus by its leaf-like bracteoles and small obtuse, mainly elliptic leaves. It flowers in December and Januaryand its fruits maturefrom April to November, being perhapsthe only species in which the fruits maturein both the fall and spring. 11. Myrceugeniaparvifolia(A. P. de Candolle)Kausel, Rev. Mirt. Chile: 2. 1940; Revista Argent. Agron. 9: 53. 1942. Figs. 10, 28D. Eugenia parvifolia A. P. de Candolle, Prodr. 3: 266. 1828.

Lumabaeckeoides Grisebach,Syst. Bemerk.:32, 1854;Abh., Kon. Ges. Wiss. Gottingen6: 120. 1856. Type. Lechler 868, "Chiloe: pr. Ancud m. Jul." (holotype, GOET, n.v.; isotype, MICH;Field Museumneg. 19918of isotype at M, photo at MICH).

Shrub or small tree 1-5 m high; hairs whitish to reddish-brown,a mixture of simple and dibrachiate; twigs light grey to reddish-brown,densely pubescent when young, the hairs spreading, deciduous with the young twig bark; leaves glabrousor with scattered symmetricdibrachiatehairs, narrowlyelliptic, oblong or lanceolate, 1-2.6 cm long, 0.3-0.7 cm wide, (2.2-)2.7-4.8 times as long as wide; apex and base acute to obtuse; midvein impressedfor entire length to not at all above, moderatelyprominentto indistinctbelow; lateral veins indistinctor up to 5 pairs faintly visible; marginal veins equalling laterals in prominence; blades subcoriaceousto submembranous,dull grey-green,often tinged with reddish-brown above, lighter grey-green or yellow-green below; peduncles uniflorous, slightly flattened, 5-14 mm long, 0.3-0.4 mm wide, glabrousor with a few scatteredhairs, solitaryin the axils of leaves; bracteoles ovate to lanceolate, 0.51 mm long, 0.2-0.5 mm wide, 1.3-3.5 times as long as wide, subcoriaceous, sparsely pubescent to glabrous within and without, loosely clasping the hypanthium; calyx-lobes ovate to oblong, 1.2-2.3 mm long, 1-2 mm wide, 1-1.7 times as long as wide, glabrous or with a few scattered hairs within and without, submembranous,concave; hypanthiumobconic, 0.8-1.5 mm long, sparsely pubescent, the hairs whitish, appressed; disk 1-2 mm across, very sparsely pubescent

Myrceugenia

to glabrous;stamens 60-100; anthers 0.2-0.3 mm long; ovary 2-3-locular; ovules 3-8 per locule; fruit globose, ca. 7 mm in diam.; seeds few, oblong, 3-4 mm long. Type. Dombey s.n., "in Peruvia"(holotype, G-DC, n.v.; possible isotype, RB; Field Museum neg. 7949 of holotype, photo at MICH). Specimensexamined. CHILE. Maule:Empedrado,29 Jan 1943, Bernaths.n. (CONC, H). Nuble: Bosque Fisco, camino Quirihuea Cobquecura,21 Mar 1971, Landrum& Donoso s.n. (EIF). Concepci6n: Laraquete,rio Cruces, 24-25 Mar 1940, Kausel 745 (F, HBR). Arauco:Contulmo, 13 Feb 1919,Behn s.n. (CONC, F). Malleco:CordilleraNahuelbuta,Dihuelhue,23-24 Feb 1939,Kausel589 (F, H, SGO), 27 Feb 1939, Kausel 614 (F, H, LP), Kausel 615 (F, H), Kausel 617 (H). Cautin: Loncoche, 15-23 Mar 1939, Bernath s.n. (CONC, F, H, HBR, SGO). Valdivia: without specific

locality, 1896, Buchtien s.n. (US), 1862, Philippi s.n. (MICH), Hohenacker 184 (MICH); San Jose

de la Mariquina,ca. 50 m, Jul 1926,Hollermayer587 (M, SI, US), Feb 1926, Hollermayers.n. (SI, US); ca. 10 km north of Valdiviaalong highwayto San Jose (ca. 39?45'S,73?15'W),ca. 100 m, 24 Feb 1978,Landrum3271 (MICH,VALD). Llanquihue:PuertoMontt, Feb 1925, ClaudeJoseph 3266 (US); Chin-Chinal N. O. de PuertoMontt, 150m, 19Feb 1942,Pisano & Geni 1273(F); las Lagunillas, 10 km W. Puerto Montt, 21 Mar 1954, Sparre & Constance 10839 (CONC, F); Est. Parga, Rio Peuchen,FundoPorvenir,22-23 Nov 1944,Kausel 1436(H). Chiloe:Ancud, nearairport,7 May 1972, Landrum888 (MICH);Castro, Piriquina[?], 150 m, Mar 1924, Werdermann299 (SI, US); Quemchi, ca. 1 km west of town (ca. 42?10'S,73030'W),near sea level, 10 Feb 1978, Landrum3188, 3192 (MICH, VALD); Yaldad, ca. 10 km west of Quell6n (ca. 43?15'S,73?45'W),near sea level, 7-8 Feb 1978, Landrum3125, 3127 (MICH, VALD); Huildad, near sea level, 8 Feb 1978, Landrum 3164 (MICH, VALD); withoutspecific locality, Feb 1925, ClaudeJoseph 3309, 3319 (US). Without specific locality: Ruiz & Pavon s.n. (F), Pavon s.n. (F), Gay s.n. (MICH).

Myrceugenia parvifolia, part of the M. ovata complex, could possibly be confused with M. ovata var. gracilis and M. leptospermoides. With the first, a

Brazilian species, it is compared at the end of Key G. Myrceugenialeptospermoides differs from M. parvifolia in having its peduncles densely pubescent and its linear to oblong leaves 1.5-3 mm wide. Myrceugeniaparvifolia is an understory shrub or tree in the wet forests of southern Chile. A map of its distributionis shown in Fig. 10. In open areas it becomes a dense shrub. It flowers mainly in Februaryand March and the fruits probablymatureduringthe springmonths of October and November. 12. Myrceugenialeptospermoides(A. P. de Candolle) Kausel, Rev. Mirt. Chile: 2. 1940;Revista Argent. Argon. 9: 52. 1942. Figs 10, 30A. Eugenia leptospermoidesA. P. de Candolle,Prodr.3: 266. 1828. Eugenia leptospermoides [var.] a microphylla Berg, Linnaea 27: 143. 1856. (Inadmissible name to be replaced by E. leptospermoides var. leptospermoides because E. leptospermoides A.

P. de Candolleis cited in synonymy).

?Eugenia leptospermoides [var.] /3 latifolia Berg, Linnaea 27: 143. 1856. Type. Philippi s.n.,

"in Chili" (holotype, B, n.v., presumablydestroyed).

Eugenia leptospermoides [var.] y longifolia Berg, Linnaea 27: 143. 1856. Type. Poeppig 421,

"in silvis Chili austr., ad Talcahuano,""MyrtusbrachyphyllaKunze in hb. Poeppig"(holotype, W). Eugenia thymifoliaR. A. Philippiex Reiche, Anales Univ. Chile 98: 719. 1897;Flora de Chile 2: 303. 1898. Type. PresumablyPhilippis.n., "Provinciade Concepci6n(PeninsulaTumbez)" (apparentholotype, SGO-64335).

Shrub0.5-2 m high; hairs reddish-brown,a mixtureof simple and dibrachiate; twigs grey to whitish-grey, densely pubescent when young, the hairs persisting until the first bark falls; leaves oblong to linear, 4-15 mm long, 1.5-3 mm wide, 2.7-7 times as long as wide, glabrousor with a few scatteredhairs beneath; apex obtuse; base obtuse to acute; midvein slightly impressed or not at all above, moderatelyprominentbelow; lateral and marginal veins indistinct; blades sub-

Flora Neotropica

coriaceous, dull grey-greenor brownish-greenabove, lighter, often yellow-green below; peduncles uniflorous, slightly flattened, 2-8 mm long, 0.2-0.3 mm wide, densely pubescent, solitary (rarely in pairs) in the axils of leaves; bracteoles ovate to oblong-lanceolate,blunt, 0.4-1.5 mm long, 0.3-0.7 mm wide, 1.3-2.3 times as long as wide, sparsely pubescent to glabrous within and without, subcoriaceous, loosely clasping the hypanthium;calyx-lobes ovate, oblong-ovateto suborbicular,1-2 mm long, 1-1.5 mm wide, 0.8-1.4 times as long as wide, glabrous to very sparsely pubescent within and without, membranousto subcoriaceous, concave, usually tinged with red; hypanthiumdensely pubescent, ca. 1 mm long, obconic to subglobose; disk 1-1.5 mm across, sparsely pubescent; stamens 60-90, ca. 3-5 mm long; anthers 0.2-0.4 mm long when dry; petals suborbicular, 1.5-2.5 mm in diam.; style ca. 4-5 mm long, glabrous to very sparsely pubescent; ovary 2-3-locular; ovules 2-6 per locule; fruit reddish-purple, globose, 4-5 mm in diam.; seeds 1-2 in fruits seen, oblong, 2-3 mm long. Type. "in Chili ex herb Deless." (G-DC, hereby designatedas lectotype; possible isolectotype, RB; Field Museum neg. 7944 of lectotype, photo at MICH); apparently Dombey s.n., "in Peruvia ex herb, Juss." (G-DC, paratype; Field Museum neg. 7944 of paratype, photo at MICH). Specimensexamined.CHILE. Concepci6n:PeninsulaTumbez, 22 Mar 1936,Pfister4810 (CONC, H), 10Mar1957,Junge3012 (US), 6 Sep 1896,Dusen s.n. (G);Talcahuano,ParqueHualp6n,(36?47'S, 73?10'W),60 m, 23 Aug 1969, Carrasco 16 (CONC); Caj6n El Cipres, 5 km al interiorde Colcura

(37?04'S, 73?03'W), 300 m, 21 Apr 1976, Marticorena et al. 1053 (CONC); Quebrada Honda, Puente

Mellizos (37?07'S,73?07'W),175 m, 21 Apr 1976, Marticorenaet al. 1070 (CONC);zwischen Santa Juanaund Coronel, Merxmiller24835(M). Arauco:Lanalhue,ClaudeJoseph 5933 (US); Lago LleuLieu, 23 Feb 1939, Kausel 602 (LP); Contulmo, Feb 1940, Grandjot3985 (CONC, H). Malleco: CordilleraNahuelbuta,Coyancahuin,400 m, 14-16 Feb 1939,Kausel551 (H); CordilleraNahuelbuta, Dihuelhue,cerca Pur6n,27 Feb 1939, Kausel 616 (SGO).

Myrceugenialeptospermoides is restricted to a small area in Chile from Concepcion to the cordilleraof Nahuelbuta.A map of its distributionis shown in Fig. 10. It is apparentlyclosely related to M. parvifolia but distinguishedfrom that species by its pubescent peduncles and its leaves which are 1-3 mm wide. In M. parvifolia the peduncles are glabrous or nearly so and the leaves are 3-7 mm wide. Myrceugenia leptospermoides is also rather similar to M. smithii of Brazil

and is comparedwith that species at the end of Key I. Myrceugenia leptospermoides flowers in February and March and the fruits seem to matureabout 1 year later. 13. MyrceugeniahatschbachiiLandrum,Brittonia32(3): 372. 1980. Figs. 10, 29D-F. Shrub about 1 m high; hairs reddish-brown,a mixtureof dibrachiateand simple; twigs reddish-brownto purple-brown,densely pubescent when young, soon glabrescent; leaves narrowly elliptic to oblanceolate, 1.5-3.7 cm long, 0.2-1.2 cm wide, 3-7 times as long as wide, glabrousor with scattered hairs; apex acute to obtuse; base cuneate; petiole slightly channeled, 1-3 mm long, 0.2-0.5 thick, pubescent when young, glabrescent with age; midvein impressed only slightly proximallyabove to not at all, moderatelyprominentbelow; lateral veins indistinct or up to ca. 10 pairs faintly visible; marginal veins equalling laterals in prominence; blades coriaceous, reddish-brown(when dry) to grey or yellowgreen above, lighter yellow-green below, the upper surface slightly lustrous to dull; peduncles uniflorous, flattened, (0.6-)1-2 cm long, 0.3-0.5 mm wide, glabrous to very sparsely pubescent, solitary in the axils of leaves; bracteoles lanceolate, 1-2 mm long, 0.5-0.8 mm wide, 2-3.5 times as long as wide, subcoriaceous, glabrous or nearly so within and without, clasping the hypanthium;

Myrceugenia

calyx-lobes ovate, 1.4-1.8 mm long and wide, ca. 1-1.2 times as long as wide, submembranous,glabrousor nearly so within and without, concave; petals suborbicular, ca. 3 mm in diameter, glabrous; hypanthiumobconic, ca. 1-1.5 mm long, glabrous to moderately pubescent; disk ca. 1-2 mm across, glabrous to sparsely pubescent; stamens ca. 80-120, 4-5 mm long; anthers ca. 0.3 mm long; style ca. 5-6 mm long, glabrous; ovary 2-3-locular; ovules ca. 3-7 per locule; fruit unknown. Type. G. Hatschbach 13043, Parana,Mun. Palmeira,Rod. do Cafe, Rio Tibagi, 22 Oct 1965(holotype, MBM; isotypes, HB, MICH, MVM). Specimens examined. BRAZIL. Parana: Limit between Mun. Ponta Grossa and Mun. Palmeira, gorge of Rio Tibagi at point where Br 376 highway crosses (ca. 25?20'S, 49?50'W), ca. 1000 m, 16 Nov 1977, Landrum 2545, 2552, 2553, 2554, 2556 (MBM, MICH); Mun. Lapa, Gruta do Monge, 22 Oct 1967, Hatschbach 17562 (MVM).

This species has been named for its originalcollector, Dr. Gert Hatschbachof the Museu Botanico Municipalof Curitiba,to whom I owe very much, for his numerouscollections and kind cooperationwhile I was in Brazil. Myrceugenia hatschbachii has been collected few times so the limits of its variation are not well understood. It is similar to two western and two eastern species with which it is comparedbelow. M. hatschbachii

M. lanceolata

Brazil Peduncles uniflorous, solitary, glabrous to very sparsely pubescent

Chile Peduncles uniflorous or bearing a 3 flowered dichasia, some usually in pairs, usually densely pubescent Calyx-lobes 1.8-4 mm long, 1-1.7 times as long as wide Hypanthium 2-5 mm long

Calyx-lobes 1.4-1.8 mm long, 1-1.2 times as long as wide Hypanthium 1-1.5 mm long M. hatschbachii

M. pinifolia

Brazil Peduncles glabrous to very sparsely pubescent, (0.6-)1-2 cm long Hypanthium 1-1.5 mm long Twigs soon glabrescent

Chile Peduncles densely pubescent, 0.5-1 cm long

Bracteoles about as long as the hypanthium

Hypanthium 1.5-2.5 mm long Twigs remaining pubescent for more than a year Bracteoles usually shorter than the hypanthium

M. hatschbachii

M. smithii

Peduncles (0.6-)1-2 cm long, glabrous to very sparsely pubescent Twigs soon glabrescent, reddish-brown to purple-brown Leaves 1.5-3.7 cm long, 0.2-1.2 cm wide, dark green above

Peduncles 0.1-0.4 cm long, densely pubescent

M. hatschbachii Leaves 3-7 times as long as wide Some peduncles usually exceeding 1.5 cm, glabrous or less often very sparsely pubescent Twigs glabrescent within about a year Shrub about 1 m high, along streams in open areas (campos)

Twigs pubescent until the first bark falls, orangish Leaves 0.8-2.5 cm long, 0.2-0.5 cm wide, light grey green above M. ovata var. gracilis Leaves mostly about 2 times as long as wide Peduncles rarely exceeding 1.5 cm, usually averaging about 1 cm long or less, usually densely pubescent but variable Twigs often remaining pubescent for over a year Shrub to small tree 2-3 m high in forests, typically along the foggy eastern edge of the planalto

Flora Neotropica

Myrceugenia hatschbachii as far as is known grows along streams in open areas in the region of campos and Araucaria forests west of Curitiba.A map of its distributionis shown in Fig. 10. It flowers mainly in October.

14. Myrceugeniasmithii Landrum,Brittonia32(3): 374. 1980. Figs. 10, 29A-C. Mvrceugenia regnelliana var. leptophylla Legrand, Darwiniana 11(2): 354. 1957, pro parte, not

as to type.

Shrub 1-3 m high; hairs reddish-brownto yellowish-brown,a mixtureof simple and dibrachiate;twigs densely pubescent when young, the hairs spreading,persisting until the first barkfalls, the older twigs often becoming smooth, orangish; leaves glabrousor with a few scattered appressed hairs, narrowlyoblanceolate, narrowlyelliptic or linear, 0.8-2.5 cm long, 2-4 mm wide, 3-7 times as long as wide; apex obtuse or bluntly acute; base cuneate, taperingfrom about midleaf; petiole slightly channeled, 1-1.5 mm long, ca. 0.5 mm thick, sparsely to densely pubescent; midvein not impressed or only slightly so proximallyabove, moderately prominentto indistinctbelow; lateral and marginal veins indistinct(rarely faintly visible); blades subcoriaceous,grey-greento brownish-greenabove, lighter yellowish-brownor grey-greenbelow, the surfaces dull; peduncles uniflorous, flattened slightly, 1.5-4 mm long, 0.2-0.4 mm wide, densely pubescent, solitary in the axils of leaves; bracteoles lanceolate, 0.9-1.5 mm long, 0.4-0.6 mm wide, 2.2-2.5 times as long as wide, submembranous,sparsely pubescent without, glabrous within, clasping the hypanthium;calyx-lobes ovate, 0.9-1.5 mm long, 0.91.2 mm wide, 0.7-1.4 times as long as wide, submembranous,sparselypubescent without, glabrouswithin, concave; petals suborbicular,ca. 2 mm in diam., brown when dry, glabrous; hypanthium obconic, ca. 1 mm long, densely pubescent; disk ca. 1 mm across, sparsely pubescent; stamens ca. 60, 2-5 mm long; anthers 0.2-0.3 mm long when dry; style 4-5 mm long, glabrous; ovary 2-3-locular; ovules 3-8 per locule; fruit unknown. Type. Smith & Reitz 12411, Mun. Indaial (Mun. Apiuna on label), marginof Rio Itajai-aqu,Subida, ca. 27?5'S,49025'W,100 m, 11 Oct 1964(holotype, HBR; isotypes, GH, MICH, NY, R, UC, US). Specimensexamined.BRAZIL.SantaCatarina:Mun. Lontras,Salto do Pilao, 350 m, 19 Oct 1958, Reitz & Klein 7368 (H, HBR); Mun. Indaial, Subida, ca. 50 m upstreamfrom bridgecrossing Rio

Itajai-aqu, 100 m, 19 Nov 1977, Landrum 2617, 2618, 2619, 2620, 2621, 2622, 2623, 2624, 2625 (MBM, MICH).

Legrand included Myrceugenia smithii in his M. regnelliana var. leptophylla,

but the type of that taxon, Reitz 2576, belongs to M. ovata var. gracilis. In the Flora IlustradaCatarinense(1970) Legrandproposed a new type, Reitz & Klein 7368, which does belong to M. smithii, but such a change in types is contraryto the rules of nomenclature.Thereforea new name has been chosen. The epithet honors Dr. Lyman B. Smithof Washington,D.C., who has made many important contributionsto Brazilianbotany throughhis numerouscollections and publications. Myrceugenia smithii appears to be derived from M. ovata var. gracilis. The

two are comparedbelow.

Myrceugenia 70 70

...

70rs'.

.40

-5''"'

. ;'... -r *:*'.*

;*~'

.,.. -'

C '

.. ..;

',.. t

,j&-'..i_.."

l ':x, '....o . ,

.,.

ZI&&M

. .SR*S

.*.'. '

"'*'*

'?. ..

0!. * '*? . *''. .a.

a a

"* . -'**-*''**'** . :*

:ii.:.-::,,' , -" v:.:-'." ,'***'--"*' V:.W,~".^A ."-^'!^ W.

FIG. 10. Distributionsof Myrceugenia ovata, M pinifolia M. obtusa, M. leptospermoides, M. parvifolia, M. hatschbachii, and M. smithii.

Flora Neotropica M. smithii

M. ovata var. gracilis

Leaves 0.8-2.5 cm long, 0.2-0.4 cm wide, 3- Leaves 1-3 cm long, 0.5-1 cm wide, 1.7-3 7 times as long as wide, nearly concolortimes as long as wide, often discolorous, the upper surfacedull or lustrous ous, the upper surfacedull Twigs often becomingorangish Twigs usually grey to reddish-brown Floweringin October Floweringmainlyfrom Novemberto January Shrubgrowingalong streamsat elevationsbe- An understoryshrubin forests mainlyabove low 400 m 800 m

Admittedly little materialhas been collected of Myrceugeniasmithii, and my collections have been of sterile plants only. Nevertheless, this species, which may be restrictedto a small geographicarea, seems to have become ecologically and phenologicallyseparatedfrom M. ovata var. gracilis and is morphologically distinguishablefrom it. Myrceugenia smithii is also similar to M. leptospermoides of Chile and is com-

pared directly with that species in Key I. It is compared with M. hatschbachii of Brazil in the discussion of that species. A distributionmap of Myrceugeniasmithii is shown in Fig. 10. 15. Myrceugeniaplanipes(Hooker et Arnott)Berg, Bot. Zeitung(Berlin) 16: 250. 1858;Linnaea 30: 670. 1861. Figs. 8K, 11, 30B. Eugeniaplanipes Hooker et Arnott, Bot Misc. 3: 323. 1833. Eugeniaplanipes [var.]a genuina Berg, Linnaea27: 162. 1856.(Inadmissiblenameto be replaced by E. planipes var. planipes.) Eugenia planipes [var.] 3 grandifolia Berg, Linnaea 27: 162. 1856. Type. Bridges 693, "in Chili'

(holotype, W). Eugenia distoma Berg, Linnaea 27: 155. 1856. Type. Chamisso 55, "Habitat in Chili" and Poep-

pig 456 "in silvis ad Talcahuano," "(v. s. in hb. Berol. et Vindob.)" (lectotype, hereby designated, Poeppig 456 at W; probableisolectotypes, Poeppig 456 at W and Poeppig 457 at P and W). Myrciaplanipes (Hooker et Arnott)Kiaerskou,Enum. Myrt. bras.: 115. 1893. Mvrceugeniadistoma (Berg) Kausel, Revista Argent. Agron. 9: 238. 1942.

Small tree up to ca. 8 m high; hairs whitish, yellowish or less often reddishbrown, symmetricallydibrachiate,appressed; twigs sparsely to densely pubescent when young, glabrescent with age, the bark grey, often persisting as strips beneath the leaf bases; leaves moderately to sparsely pubescent below when young, glabrescentwith age, elliptic, 2.2-8 cm long, 1-3 cm wide, 2-3.4 times as long as wide; apex acuminate,less often acute; base cuneate to acuminate;petiole channeled, 2-6 mm long, 1-2 mm thick, densely to sparsely pubescent, becoming less so with age; midvein impressedfor entire length or nearly so above, prominent below; lateral veins indistinguishableor up to ca. 30 pairs faintly visible; marginal veins equalling laterals in prominence; blades drying light to dark grey-greenor reddish-brownabove, lighter yellow-green, grey-greenor tan beneath, coriaceous;peduncles stronglyflattened,0.7-3 cm long, ca. 1 mm wide, sparsely to moderatelypubescent, solitary or up to 3(-4) in the axils of leaves or less often in the axils of bracts; bracteoles ovate to lanceolate, 1.9-3.1 mm long, 0.8-1.3(-1.7) mm wide, 1.4-3 times as long as wide, sparselypubescentwithinand without, clasping the hypanthium or somewhat reflexed; calyx-lobes broadly ovate, concave, 1.8-3 mm long, 2.5-4 mm wide, 0.5-0.8(-1.2) times as long as wide, coriaceous, sparselypubescentto glabrouswithinand without;hypanthium hemiobicular,densely pubescent, 1.5-4 mm long, usually prolonged 1-1.5 mm past the ovary; disk 2.5-4 mm across, sparsely pubescent to glabrous; style ca. 1 cm long, glabrous; stamens 120-220, 7-12 mm long; anthers 0.4-0.8 mm long when dry; petals suborbicular,3-6 mm in diam.; ovary 3-4-locular; ovules 411(-13) per locule; fruit globose, 0.8-1 cm in diam., purplish-black;seeds few, ca. 6 mm in diam.

Myrceugenia

Type. Cuming28 "Chiloe" (holotype, E-GL, n.v.; photo of holotype at A, negative at SGO); probable isotype, Cuming 98 at W; Field Museum neg. 31599 of Cuming 98 at W, photo at MICH).

Specimens examined. CHILE. Concepci6n:Ramuncho, 14 Dec 1941, Behn s.n. (CONC, H), 22 Dec 1946,Behn s.n. (H); ParqueHualpen,ca. 15km west of Concepci6n(ca. 36?50'S,73?10'W),near sea level, 29 Jan 1978, Landrum3106 (MICH, VALD); Concepci6n, 1896, Neger s.n. (C, W); Laraquete, quebradasobre la estaci6n, 20-60 m, 14-27 Feb 1949, Kausel 2795, 2796 (H). Arauco: Canete, Nov 1927, Claude Joseph 5362 (US); Curanilahue, Nov 1927, Claude Joseph 5396 (US);

Lanalhue,Jan 1929, Claude Joseph 5936 (US); Contulmo, 11 Jan 1919, Behn s.n. (F); Antiquina, Lago Lleu-Lleu, 23-24 Feb 1939, Kausel 599 (F), Kausel 596 (H); Isla Mocha, 270 m, Kunkels.n. (H). Cautin:Pitrufqu6n,Rio Pucul6n (Tolten), 21 Feb 1942, Kausel 1069 (H, HBR, LP). Valdivia: San Jose de la Mariquina,25 Oct 1939,Aravena 18057(UC), ca. 50 m, Jan 1926, Hollermaver1207 (MO, NY, SI, US), Jul 1925, Hollermayer1208 (MO, NY, SI, US); Corral, 20 Jan 1935, 25 m, Gunckel 5054 (GH, W), 8 Feb 1935, Gunckel 1476.4 (GH), 21 Jan 1906, Sargent s.n. (A), 1905-1906,

Thaxters.n. (GH), 50 m, 24 Dec 1935, West4861 (GH, MO, UC, US), 400 m, 18 Apr 1936, Mexia 8023 (F, GH, UC, US); Niebla, 3 Jan 1933,Junge 852 (F); Los Lagos, 24 Jan 1931,Behn 1175(MO, SI); Puerto Nuevo, 11 Feb 1972, Cantino80 (GH); San Juan, 8 Mar 1935, Gunckel88 (F); Coreltue [?], 10 Mar 1923, Gunckel2131 (SI); Valdivia,Amargo, 14 Jan 1934, Montero1335 (GH); Valdivia, 1888, Philippi s.n. (US), 29 Jan 1897, Buchtiens.n. (GH, US). Osorno: 10 miles east of ocean on road to termas de Pucatrihue,29 Jan 1958, Eyerdam10528a(UC, US); Volcan Antillanca,ca. 1000 m, 28 Jan 1958, Everdam 10519 (F, NY, UC); Puyehue, LagunaEspejo, ca. 400 m, 20 Feb 1964, Kausel 4799 (H). Llanquihue:Puerto Montt, 24 Jan 1909, Hicken s.n. (SI); Cayetue, Feb 1927, Hicken s.n. (SI); Pargua,18 Feb 1964, Kausel4760 (H); Seno Reloncavi,Isla Huar(41?40'S),12 Mar 1952,Poulsen s.n. (C). Chiloe:withoutspecificlocality, Feb 1925,ClaudeJoseph 3304 (US); Ancud, Lechler875 (F), 14 Nov 1968, Cunninghams.n. (GH); Ancud, near airport,7 May 1972, Landrum 947 (EIF, MICH);Quemchi, ca. 1 km west of town (ca. 42?10'S,73?30'W),near sea level, 10 Feb 1978, Landrum3179 (MICH, VALD); Castro, ca. 14 km north on road to Ancud (ca. 42?20'S, 73?40'W),9 Feb 1978, Landrum3177, 3178 (MICH, VALD); Yaldad,ca. 10 km west of Quell6n(ca. 43?15'S, 73?45'W), 1-5 m, 7-8 Feb 1978, Landrum 3123, 3143 (MICH, VALD); Huildad, near sea

level, 8 Feb 1978, Landrum3165 (MICH, VALD). Aisen: Aisen, 16-17 Nov 1944, Grosse 77 (H); Ais6n, PuertoPuyuhuapi,ca. 10 m, 30 Oct 1939,Schwabe 10 (NY); 23 Dec 1939, Schwabe lOa (NY); Port Otway, 9 Feb 1888, Lee s.n. (US); Rio Exploradores,entre EstanciaTeresa y Bahia Exploradores (46?25'S,73?10'W),40 m, 3 Mar 1967, Seki 350 (CONC).Provinceunknown:Mal Paso, 8 Feb 1930, Gunckels.n. (MO); Quitaluto,23 Mar 1935, Gunckel1453.4 (GH); CordilleraChaihuin,3 Jan 1932,Gunckel3062 (GH); Cordillerade Alerce, Mar 1969,Reed s.n. (GH);Antilhue[?], 23 Jan 1906, Sargent s.n. (A); without specific locality, Gay s.n. (GH, MICH). ARGENTINA. Neuquen: Quetrihue,pr6ximo a la punta, 4 Feb 1959, Boelcke 10548 (CTES); Quetrihue,8 Mar 1942, Diem s.n. (H).

At W there are two specimens of this species labeled Cuming98. They fit the description of Eugenia planipes of Hooker and Arnott well and the 9's could easily become 2's with only slight modification.I therefore suspect that they are isotypes. ApparentlyPoeppig's numbers456 and 457 have been mixed together. One of the collections must have been Myrceugeniaplanipes while the other was Luma

apiculata. At W there is a sheet each of Poeppig 456 and 457 which are Luma apiculata and also three sheets of Poeppig 456 and one of Poeppig 457 which are M. planipes. Two of the sheets of Poeppig 456 which are M. planipes have

"Eugenia distoma Bg." written on them by Berg himself. One of these has been selected as the lectotype.

Myrceugenia planipes is apparently most closely related to M. reitzii and M. kleinii of Brazil and is compared with them in Key A. Myrceugenia correifolia

of Chile may also be a close relative as is apparentfrom various floral characteristics but it has leaves which are quite differentand is geographicallyseparated

from M. planipes. Myrceugenia chrysocarpa is the species which might most

easily be confused with M. planipes because they grow more or less sympatrically. The two are comparedin the discussion of M. chrvsocarpa and at the end of Key M. Myrceugenia planipes is an understory tree in the rain forests of southern

Flora Neotropica

Chile, growing mainly at elevations below 700 m. In Argentinait has been found only on the QuetrihuePeninsula, which seems to be a locality of relatively moderate temperaturesand heavy rain fall in Lago Nahuel Huapi. A map of its distributionis shown in Fig. 11. 16. Myrceugenia kleinii Legrand et Kausel, in Legrand, Sellowia 13: 306. 1961. Figs. 11, 30D. Tree 6-18 m high; hairs whitish or yellowish, more or less appressed, dibrachiate, mainlysymmetrical;twigs moderatelyto sparselypubescent when young, glabrescentwith age, stronglyflattenedat the nodes at first, becoming terete, the bark smooth whitish-grey; leaves elliptic to obovate or oblanceolate, 4-11 cm long, 1-4 cm wide, 2-4 times as long as wide, glabrous or very sparsely pubescent; apex acute to acuminate;base cuneate to acuminate;petiole 6-15 mm long, 1-1.5 mm thick, shallowly channeled, sparsely pubescent to glabrous; midvein deeply impressed proximallyto shallowly or not at all impressed distally above, prominent below; lateral veins indistinguishableor up to ca. 20 pairs barely visible; marginalveins equallinglateralsin prominence;blades dryingdarkgreygreen or lightergrey-greenor yellow-greenabove, aboutthe same color but lighter below, coriaceous;flowers mainly aggregatedtogether in bracteate shoots, with up to ca. 7 flowers, the bracteate shoots ca. 1.5-4 cm long, solitary or 2 or 3 in a row in the axils of leaves, terminatingin a flower; peduncles lacking to 15 mm long, 0.5-1 mm wide, stronglyflattened, sparsely to moderatelypubescent, usually solitaryin the axils of bracts or rarelyin the axils of leaves; bracts lanceolate to linear, 2-5 mm long, moderately to densely pubescent; bracteoles ovate to lanceolate, 2-3.7 mm long, 0.8-1.1 mm wide, 1.8-3.7 times as long as wide, sparsely pubescent to glabrous within, sparsely to densely pubescent without, clasping the hypanthium;calyx-lobes broadly ovate, concave, 1.8-3 mm long, 1.9-3.2 mm wide, 0.7-1 times as long as wide, coriaceous, sparsely pubescent without, sparsely pubescent to glabrous within; hypanthium obconic, densely pubescent, 2-3 mm long; disk ca. 3 mm across, very sparsely pubescent to glabrous; style 5-7 mm long, glabrous; stamens 130-170, 5-9 mm long; anthers 0.5-

0.8 mm long when dry; petals suborbicularto slightly broaderthan long, 2.5-5 mm in diam.; ovary 3-4-locular; ovules 4-10 per locule; fruit unknown. Type. Klein 1540, Santa Catarina,Itajai, Cunhas (holotype, MVM; isotypes, F, H, HBR, UC, US). Specimens examined. BRAZIL. Parana:Mun. Guaratuba,Pedra Branca de Araraquara,24 Sep 1964,Hatschbach 11578(MBM, MICH,MVM);Mun.Guaraque9aba,9 Aug 1967,Hatschbach16901 (C, HB, MBM, MVM, MO, UC, US). Santa Catarina:Mun. Itajai, Morroda Ressacada, 250 m, 15 Sep 1955, Klein 1579 (H, HBR, MVM, NY, UC, US).

Myrceugeniakleinii is known from only four collections but it is a distinctive species. Morphologicallyit falls about midway between M. reitzii and M. planipes and it is comparedwith these in Key A. Myrceugeniakleinii seems to be confinedto the lowlandcoastal regionof Santa Catarinaand Parana. A map of its distributionis shown in Fig. 11. It flowers in the late winter and springmonths of August and September. 17. Myrceugenia reitzii Legrand et Kausel, in Legrand, Sellowia 13: 305. 1961. Figs. 11, 30C. Tree or shrub 3-10 m high; hairs whitish, yellowish, dibrachiate,usually symmetric or nearly so, usually appressed; twigs densely pubescent when young,

Myrceugenia

glabrescentwith age, stronglyflattenedat the nodes at first, becomingterete with age, the bark light whitish-greyto reddish-tan;leaves moderatelypubescent beneath when young, becoming less so or glabrescent with age, elliptic, oval or oblong, 4.5-19 cm long, 2.3-8.5 cm wide, 1.8-2.8 times as long as wide; apex broadly rounded, acute or abruptlyacuminate; base rounded to cuneate, sometimes oblique;petiole 0.8-3.5 cm long, 1-2.5 mm thick, pubescent when young, glabrescentwith age, channeleddistally; midvein impressedproximallyto not at all distally above, prominentbelow; lateral veins indistinguishableor up to about 20 pairs moderatelyprominent;marginal veins equallinglaterals in prominence; blades drying yellow-green, grey-greento reddish-brownabove, lighter yellowgreen or grey-green below, coriaceous; peduncles strongly flattened, 1-2.5 cm long, 1.5-2 mm wide, densely pubescent, solitary or in pairs in the axils of leaves or bracts;flowers normallyborne on bracteate shoots up to ca. 7 cm long, these terminalor axillary, usually with not more than 8 flowers, terminatingin a vegetative bud; bracts 3-5 mm long, densely pubescent, usually deciduous at about anthesis; bracteoles lanceolate, 5-7 mm long, 1.7-3.2 mm wide, 2-3.4 times as long as wide, sparsely to moderately pubescent within, moderately to densely pubescent without, subcoriaceous,clasping the hypanthium;calyx-lobes broadly ovate to hemi-orbicular,3.5-6 mm long, 4-8 mm wide, 0.6-1 times as long as wide, concave, densely pubescent without, densely to sparsely pubescent within; hypanthium obconic, densely pubescent, 3-5 mm long; disk 3-6 mm across, sparsely pubescent to glabrous; style ca. 7-8 mm long, very sparsely pubescent to glabrous;stamens 300-400, 5-15 mm long; anthers 0.6-1 mm long when dry; petals suborbicular,ca. 5-10 mm in diam.; ovary 3-4-locular; ovules (5-)8-17 per locule; fruit globose, 1-1.5 cm in diam., grey to grey-brownwhen dry; seeds few, 1-3 in fruits seen, ca. 5-7 mm long. Type. Klein 1606, Santa Catarina,Itajai, Cuhnas (holotype, MVM; isotypes, H, HBR, UC, US). Specimensexamined.BRAZIL.Parana:Mun.Antonina,Estr. Cacatu-SerraNegra, Rio Cachoeira, 17 Sep 1965,Hatschbach 12797(MBM,MVM,US); Mun. Bocaiuvado Sul, Sesmarias,Rio Capivari, 11 Nov 1968,Hatschbach20238(MBM, UC); P6rtodo Cima, 12 Nov 1910,Dusen 10263(MO);Mun. Guaraque;aba,Fdr. Abobreira,20 m, 15 Oct 1969, Hatchbach 22456 (C, CTES, MBM, MO, NY, UC); Mun. Guaraque9aba,Ribeiraodo Bananal,7 Oct 1970, Hatschbach 24879 (MBM, RV, UC); Mun. Guaraque9aba,Serrinha, 100 m, 13 Sep 1967, Hatschbach 17182 (MICH, MO, MVM, UC); Mun. Guaratuba,PedraBrancade Araraquara,12 Sep 1962, Hatschbach9243 (HBR, MBM, MICH, MVM); Mun. Morretes,Rio Mae Catira, 12 Oct 1972, Hatschbach30532 (MBM, NY); Mun. Morretes, Estradada Graciosa, Ferradura,29 Sep 1962, Hatschbach 9258 (HB, MICH, MVM); Mun. Morretes, Via Graciosa, Grota Funda (ca. 25?20'S,48?50'W),13 Dec 1977, Landrum2872, 2876 (MVM, MICH). Santa Catarina:Mun. Brusque, S. Pedro, 50 m, 13 Jan 1950, Klein 72 (H, HBR); Mun. CampoAlegre, Sao Miguel, 850 m, 24 Sep 1960, Cabrera,Reitz & Klein 9973 (H, HB, HBR, MBM, NY, UC, US); Mun. Campo Alegre-Joinvile,crest of Serra do Mar east of Campo Alegre, 900-1100 m, 10 Mar 1957, Smith & Klein 12006 (HBR, MICH, US); Mun. Itajai, Cunhas, 10 m, 29 Oct 1954, Klein 828 (H, HBR, NY), 29 Sep 1955, Klein 1610 (F, H, HBR), 5 Aug 1954, Reitz &

Klein 2028 (HBR, UC), 15 m, 26 Aug 1955, Klein 1546 (H, HBR), 24 Sep 1956, Reitz & Klein 2122 (F, H, HBR, MBM).

Myrceugenia reitzii may be most closely related to M. kleinii and grows sym-

patricallywith that species in the coastal lowlands of Santa Catarinaand Parana. It seems to be more common than M. kleinii and unlike M. kleinii, it is known to ascend to elevations of ca. 1000m in the Serrado Mar. Perhaps M. kleinii will be found at these higherelevations when more collections are made. Myrceugenia kleinii can easily be distinguishedfrom M. reitzii by its calyx-lobes which are only 1.8-3 mm long, and 1.9-3.2 mm wide and which are sparsely pubescent without.

Flora Neotropica

60 70...

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30 i;

.'.

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:%-D

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Dist.,butions of M..rceugenia correifolia, M. planipes, M. rezi,

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and M

lein.

Myrceugenia

18. Myrceugenia correifolia (Hooker et Arnott) Berg, Linnaea 30: 670. 1861. Fig. 11. Eugenia correaefolia Hooker et Arnott, Bot. Misc. 3: 319. 1833. Eugenia maritima Barneoud, in Gay, Fl. chil. 2: 391. 1847. Type. Gay s.n., "provincia de Colchagua" (holotype, P, n.v.; Field Museum neg. 36981 of holotype, photo at MICH). Luma correaefolia (Hooker et Arnott) Gray, U.S. Expl. Exped., Phan.: 542. 1854. Eugenia thalassaia Berg, Linnaea 27: 179. 1856. New name for Eugenia maritima Bameoud. Myrceugenia johowi Gusinde, Anales Univ. Chile 104: 307. 1917. No type designated or specimens cited. Myrceugenia thalassaia (Berg) Gusinde ex Fuentes, Bol. Mus. Nac. Hist. Nat. (Chile) 13: 108. 1930. Myrceugenia maritima (Barneoud) Kausel, Rev. Mirt. Chile: 2. 1940.

Tree or shrubup to 4 m tall or perhapstaller; hairs reddish-brownto whitishgrey, appressed, dibrachiate, about symmetric; twigs densely pubescent when young, glabrescent with age; leaves sparsely to densely pubescent below, glabrous to densely pubescent above, oval to elliptic, 2-7.5 cm long, 1-4.5 cm wide, 1.5-2.3 times as long as wide, the marginsrevolute, the whole leaf sometimes curling in dried specimens; apex broadly rounded or obtuse, less often acute; base roundedto broadlycuneate;petiole densely to sparsely pubescent, 2-5 mm long, 1-1.5 mm thick, scarcely channeled; midvein slightly recessed proximally to not at all distally, more pubescent than surroundingblade above, prominent below; lateral veins indistinguishableor up to ca. 10pairsbarelyvisible; marginal veins equallinglaterals;blades dull, light yellow-greento lightbluish-greenabove, grey-green to yellow-green below, coriaceous; peduncles uniflorous, flattened, 1-3.5 cm long, 1-1.5 mm wide, densely pubescent, solitary or in pairs in the leaf axils; bracteoles lanceolate, 3-7 mm long, 1-2 mm wide, 2.5-3.5 times as long as wide, coriaceous, glabrous to densely pubescent within, sparsely to densely pubescent without, clasping the hypanthium,becoming reflexed distally with age (rarely leaf-like and not clasping the hypanthium);calyx-lobes concave, coriaceous, very sparsely pubescent to glabrous within and without, conspicuously glandular, 2.5-4 mm long, 2.9-4.5 mm wide, 0.7-1 times as long as wide; hypanthium obconic, 2.5-4 mm long, densely pubescent; disk 3-4 mm across, sparsely pubescent; style 0.5-1 cm long, glabrous; stamens 130-200, 4-8 mm long; anthers 0.5-0.9 mm long when dry; petals suborbicular,concave, ca. 4-6 mm in diam., white to cream colored; ovary 3-4-locular, the ovary wall thick, glandular;ovules 6-12 per locue; fruit dark amber-brownto black, glandular, 0.7-1 cm long, 0.4-0.8 cm wide, sparsely puberulent;seeds usually 3-4, 6-8 mm long, more or less oblong. Type. Cuming 95 "Chili" (syntype, probably K, n.v.; Field Museum neg. 19930 of isosyntype at M, photo at MICH); Cruckshankss.n., "Valparaiso" (syntype, probablyK, n.v.). CHILE. Coquimbo: Fray Jorge, 680 m, 23 Jan 1942, Wagenknecht s.n. (H, LIL); 650 m, 15 Aug 1917, Skottsberg 816 (NY); 500 m, Nov 1925, Werdermann 900 (SI, US), 4 Nov 1970, Simon 233 (MICH); Cerro Talinay, 600-620 m, 8 Dec 1953, Kausel 3811 (H); Los Vilos, Jan 1899, Reiche s.n. (SGO), 7 Sep 1958, Schlegel & Consigny s.n. (EFI). Aconcagua: Zapallar, Quebrada del Tigre, 27 Feb 1952, Boelcke 6467 (MO), Boelcke 6582 (F, MO); Zapallar, Co. Frances, 650 m, 4 Jan 1949, Boelcke 3979, 3983, 3985 (F). Valparaiso: Valparaiso, 1838-1842, U.S. Exploring Expedition under Capt. Wilkes s.n. (US, possible duplicates GH, NY, PHIL); Fundo San Juan entre Quintay y Placilla, 4 Feb 1941, Kausel 920, 926 (H), Kausel 925 (H, HBR, LP), Grisai [?] s.n. (CONC, H, HBR, LP); Mirasol, ca. 36 km north of San Antonio (ca. 33?20'S, 71?40'W), 20-30 m, 26 Mar 1978, Landrum 3423, 3436, 3439 (MICH, SGO); Mirasol, 25 Feb 1955, Kausel 4160 (F, H), 14 Feb 1959, Kausel 4561 (H), 4 Mar 1952, Boelcke 6517 (CTES, F, MO). Santiago: El Tabo, Oct 1964, Kausel 4911 (H); Cartagena, Jan 1920, Claude Joseph 953 (US). Colchagua: Philippi 667 (SGO). Curic6: Lago Vichuquen, Punta Totorilla, 6 Feb 1969, Villagrdn & Tapia s.n. (SGO).

Flora Neotropica

This species is easily distinguished from all other species of Myrceugenia because of its combination of large flowers, large leaves and revolute leaf margins. It may be most closely related to M. planipes, M. reitzii and M. kleinii, being quite similar to them in the distribution of pubescence on the flowers and in the size and shape of the floral structures. None of these species has the strongly revolute leaf margins of M. correifolia nor is any as densely covered with hairs on the lower leaf surface. Myrceugenia correifolia seems to follow closely the often foggy coast of central Chile, where it apparently finds the proper combination of humidity and warm temperatures. A map of its distribution is shown in Fig. 11. It flowers from August to February and matures its fruits within a few months. 19. Myrceugenia chrysocarpa (Berg) Kausel, Rev. Mirt. Chile: 2. 1940; Revista Argent. Agron. 9: 58. 1942. Figs. 14, 31A. Eugenia chrysocarpa Berg, Linnaea 27: 168. 1856.

EugeniaphilippiiBerg, Linnaea27: 145. 1856.Type. Philippis.n., "Habitatin Chili ad radicem montis, flammaseructantis,Pise, altitudine2000', floret Martio"(holotype, W). Eugenia buxifoliaR. A. Philippi,Linnaea28: 640. 1857.Type. Philippis.n., "In radicevulcani Pise vulgo de Osornodicti Febr. 1852legi." (holotype, SGO, n.v.). Eugenia patagonica R. A. Philippi, Linnaea 33: 72. 1864. Type. Cox s.n., "Ad lacum Nahuel-

huapiinvenit orn. Cox" (lectotype, hereby designatedSGO-064321). Eugeniapetiolata R. A. Philippi,Anales Univ. Chile 84: 757. 1893.Type. PresumablyPhilippi s.n., "Habitatin valle fluminisPalena"(probableholotype, SGO-064322). Mvrceugeniabuxifolia(R. A. Philippi)Reiche, Anales Univ. Chile98: 712. 1897;Fl. Chile2: 296. 1898. LuLmachrysocarpa(Berg) BurretNotizbl. Bot. Gart. Berlin-Dahlem15: 524. 1941. Lutmaphilippii (Berg) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 523. 1941.

Shrub up to about 3 m high, without glands or at least not conspicuously glandular; hairs reddish-brown to whitish, mainly appressed, dibrachiate; twigs moderately pubescent to glabrous when young, soon glabrescent; leaves glabrous or with a few scattered, inconspicuous hairs, usually elliptic, ovate or lanceolate, less often obovate or oblanceolate, 1.3-5(-6) cm long, 0.5-2.5 cm wide, 1.5-4.5 (-5.8) times as long as wide; apex acute, obtuse or rounded; base obtuse, acute or cuneate; petiole channeled, 3-6 mm long, 0.5-1 mm thick, glabrous to very sparsely pubescent; midvein impressed for entire length or nearly so above, moderately prominent below, at least proximally; lateral veins indistinct or up to about 15 pairs faintly visible; marginal veins equalling laterals in prominence; blades coriaceous, light grey-green, yellow-green to somewhat olive-green above, light grey-green or yellow-green below, often concolorous, both surfaces dull; peduncles uniflorous, flattened, 3-16 mm long, 0.6-1 mm wide, glabrous to sparsely pubescent, solitary or in pairs in the leaf axils; bracteoles lanceolate, 1.3-2.7 mm long, 0.6-1.4 mm wide, 1.9-2.7 times as long as wide, coriaceous, clasping the hypanthium, glabrous within, glabrous or with scattered hairs without; calyx-lobes ovate to ovate-oblong, 1.7-3.2 mm long, 1.5-3.4 mm wide, 0.81.4 times as long as wide, subcoriaceous, glabrous to very inconspicuously and sparsely pubescent within and without, concave; petals glabrous, white to tan when dry, suborbicular, 2-4 mm in diam.; hypanthium obconic, the walls often slightly concave, 1.3-2 mm long, densely pubescent to nearly glabrous; disk 23 mm across, glabrous or with occasional hairs; stamens 70-150, 3-10 mm long; anthers slightly elongate, 0.3-0.5 mm long; style glabrous, 7-10 mm long; ovary 2-3-locular; ovules 3-10 per locule; fruit globose, 7-12 mm in diam., orangebrown (to purple?); seeds 1-5 in fruits seen, oblong, ca. 4-5 mm long. Type. Poeppig 926, "Habitat in Chili australis silvis imperviis ad Valle de

Myrceugenia

Quillay Leucu montiumAndes de Antuco," "v. in hb. Vindob. et Berol." (Field Museum neg. 31563 of syntype at W, photo at MICH). CHILE. Malleco: Baniosde Tolhuaca, 20 Mar 1933, Looser 2732 (F), 17 Jan 1906, Sargent s.n. (A); Bafios de Tolhuaca, ca. 1000 m, 28 Feb 1947, Kausel 2441 (HBR, LIL), 1000-1180m, 9 Mar 1939, Morrison & Wagenknecht 17449 (UC), 1200 m, 20 Feb 1935, Montero 2206 (GH); Baios de

Tolhuaca,Salto de la Culebra,28 Feb 1947, Kausel 2434 (H); LagunaVerde, 1250m, 28 Feb 1947, Kausel 2419 (CONC, F, H, HBR); Cordillerade Nahuelbuta,Coyancahuin,600 m, 14-16 Feb 1939, Kausel 537 (F), Kausel 540 (H, SGO), Kausel 545 (LP), Kausel 546, 535 (CONC). Cautin: Volcan Llaima, 1250 m, 14 Mar 1954, Sparre & Constance 10677 (UC), ca. 1200 m, Feb 1927, Werdermann

1258 (GH, SI, UC, US), ca. 1100 m, Feb 1927, Werdermann1248 (GH, SI, UC, US). Valdivia: Rifiihue, 1000m, 17 Feb 1941, Kausel 934 (H), ca. 900 m, 17 Feb 1941, Kausel 933 (F); Cordillera Pelada, 2 Nov 1965, Ricardi, Marticorena& Matthei 1214 (CONC);Reserva Forestal Llancacura, caminoHuenchucona,11 Mar 1955, Consignys.n. (EFI); Llifen, Cerrillos,25 Feb 1958, Marticorena & Furet 83 (CONC). Osorno:summitof Cordillerade la Carpa,ca. 950 m, 1-3 Feb 1958, Eyerdam 10551(F, NY, UC); Refugio La Picada, Volcan Osorno, 1 Feb 1937, Rudolph4578 (VALD). Llanquihue:Alerzal Candelaria,21 Jan 1944, Bernath886 (F), Bernath871 (CONC). Chiloe:Trumao,28 May 1932, Junge s.n. (H).

ARGENTINA.Neuquen:ParqueNacional Lanin, Lago Huechulafquen,16 Feb 1974,Correa5610 (MICH);ParqueNacional Lanin, Lag. Verde al S del Epulafquen,4 Mar 1968,Eskuche& Klein0282 (CTES);Lago Nahuel Huapi, Rinc6n,9 Feb 1940,Diem 106 (LP). Neuquen-RioNegro: Lago Nahuel Huapi, Puerto Blest (ca. 41?S, 71?50'W), ca. 800 m, 9 Feb 1938, Ferrovia s.n. (LIL), 14 Dec 1944,

Descole 2599 (LIL), 5 Mar 1946,Barba 1003(LIL), 19 Dec 1935, West4831 (GH, UC), 30 Nov 1937, Kalela 1165, 1191 (H), 5 Mar 1977, Landrum 3341, 3342, 3343, 3347 (LP, MICH); Lago Los Cantaros

(Puerto Blest), 4 Mar 1946, Barba 1052 (W). Rio Negro: Lago Frias, Dec 1944, Meyer s.n. (LIL), Meyer 7363 (W), 1 Dec 1937, Kalela 1228 (H), 25 Feb 1952, Pedersen 1607 (C), Hunnewell 16062

(GH); ParqueNacional Nahuel Huapi, Pauquaco, 16 Mar 1948, Barba 2283 (LIL); ParqueNacional Nahuel Huapi, Lago Roca, 20 Jan 1952, Boelcke & Correa5834 (CTES, SI); Lago Nahuel Huapi, Brazo Tristeza, 2 Nov 1940, Diem 93 (LP). Chubut:Lago Men6ndez, brazo norte, 2 Feb 1945, Castellanos s.n. (F, LIL).

There is a specimen at SGO (Krause s.n., "Corral1861")identifiedby Philippi

as Eugenia buxifolia, which is Myrceugenia chrysocarpa. It is therefore probable that Philippi's E. buxifolia is a synonym of M. chrysocarpa. Myrceugenia chrysocarpa is most similar to M. glaucescens of eastern South

America and M. planipes of western South America. The three are comparedat the end of Key M, but it may be emphasized here that M. chrysocarpa differs from M. planipes in the following ways: M. chrysocarpamainlygrows above 700 m elevation whereas M. planipes mainly grows below 700 m; M. chrysocarpa usually has few or no glands, whereas M. planipes is usually noticeablyglandular; M. chrysocarpararelyhas leaves over 5 cm long, the apex being acute or obtuse, whereas M. planipes has leaves often surpassing5 cm with an acuminateapex; and M. chrysocarpa is usually glabrous or nearly so except for the hypanthium whereas M. planipes often has densely pubescent peduncles and twigs. Myrceugenia chrysocarpa, probably the most cold-tolerant of the western species, grows mainly in forests of Nothofagus dombeyi at elevations above 700 m. A map of its distributionis shown in Fig. 14. It flowers in Februaryand March and its fruits maturefrom November to February. 20. Myrceugeniaglaucescens(Cambessedes)Legrandet Kausel, in Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo 1(7): 7. 1943. Eugenia glaucescens Cambessedes,in Saint-Hilaire,Florabrasiliaemeridionalis2: 368. 1833.

Shrub or small tree 2-10 m high; hairs reddish-brownto whitish, appressed, more or less symmetrically dibrachiate; twigs glabrous to sparsely pubescent when young, soon glabrescent;leaves narrowlyelliptic, obovate or oblanceolate, (2-)2.5-8 cm long, 0.5-3 cm wide, (1.5-)1.8-5(-8) times as long as wide, glabrous or with a few scatteredhairs; apex acuminate,acute to rounded;base acuminate,

Flora Neotropica

cuneate or acute; petiole channeled, glabrousto moderatelypubescent, 2-8 mm long, 0.5-1 mm thick; midvein impressed for entire length, only proximally or not at all above, prominentbelow; lateral veins indistinct or up to ca. 20 pairs barely visible; marginalveins equallinglateralsin prominence;blades grey-green to reddish-brownabove, lighter grey-green, yellow-green to light yellow below, coriaceous to submembranous,the upper surface dull to slightly lustrous; peduncles uniflorous, flattened, 1-3 cm long, 0.5-1 mm wide, glabrous to moderately pubescent, solitary or in pairs in the leaf axils; bracteoles ovate to lanceolate, (0.9-)1.3-2(-2.4) mm long, (0.5-)0.8-1.6(-2) mm wide, 1.2-2 times as long as wide, glabrousto very sparsely pubescent within and without, subcoriaceous, claspingthe hypanthium;calyx-lobes more or less hemiorbicular,1.6-3 mm long, 1.2-3.5 mm wide, 0.8-1.4 times as long as wide, densely pubescent to glabrous within, sparsely pubescent to glabrous without, subcoriaceous to coriaceous, concave; hypanthiumdensely pubescent to glabrous, 1-3 mm long, obconic, the sides sometimes slightly concave; disk 2-3 mm across, glabrousto sparsely pubescent; style 6-10 mm long, glabrous to sparsely pubescent; stamens 140-250 (-270), 3-10 mm long; anthers 0.3-0.6 mm long when dry; petals suborbicular, concave, 3-5 mm in diam.; ovary 2-4-locular; ovules (4-)6-9(-13) per locule; fruit globose to slightly elongate, 5-12 mm in diam., dark reddish-brownto purple; seeds 1-5 per fruit as far as known, oblong, 4-6 mm long. Of evolutionary interest but of taxonomic annoyance is the existence of a continuumof forms that includes Myrceugeniaglaucescens and the eastern population of M. ovata: M. ovata var. gracilis and M. ovata var. acutata. The ends

of the continuum(excluding the western populationof M. ovata) are M. ovata

var. gracilis and M. glaucescens var. glaucescens

which are quite different,

grow sympatrically,and show no signs of hybridization.But M. ovata var. acutata and M. glaucescens var. latior seem to merge with one another, as is shown in the scatter diagramin Fig. 12. Each point represents one specimen and the numericalvalues are averages of a few measurements.The points can be divided almost exactly accordingto limits of the taxa recognized here, section A corre-

sponding to M. ovata var. gracilis, B to M. ovata var. acutata, C to M. glaucescens var. latior and D to M. glaucescens var. glaucescens. For each group

the average stamen number, locule number, and ovule numberhas been calculated and these are given in Table V. Clearly there are no sharp breaks in the charactersanalyzed here. There are three possible explanationsfor the continuousnatureof the variation in this group. First, the complex could be a phylogeneticcontinuum,i.e., a dine that has turnedback on itself so that the ends are growingtogether and acting as separate species (Fig. 13A). Second, parallel evolution could have given rise independentlyto two forms that are so alike that they are inseparable(Fig. 13B). Third,two differentphylogeneticlines could have mergedto form a hybridwhich links them together (Fig. 13C). Both Myrceugenia glaucescens and M. ovata

have apparentlyclose relatives that are includedin the diagramsillustratingthese hypotheses (Fig. 13). If the first hypothesis is accepted then it is necessary that the continuumin eastern South America must be ancestral to one or both of the groups of related species. But the continuousnatureof their variationwould indicatethat members of the continuumhave only begun to diverge from each other, an unlikely situation for an ancestralgroup. Such extraordinaryparallelevolution as the second hypothesis entails does not seem likely either. Because hybridizationis known to occur at present between other species I suggest that the third hypothesis is the most probable, but the question has by no means been answered with cer-

1*. . ..

. * -., .

. '^ * .** :'... .'****:

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;

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.-......-.... **. ....

**** . * .* **** :. -

N~~~C ' :"

........

FIG. 12. Scatter diagram of Myrceugenia glaucescens (white dots) and eastern population of M. ovata (b B = M. ovata var. acutitat; C = M. glaucescens var. ltior; D = M. glaucescens var. glaucesens. See text f

Flora Neotropica

66 PL..

.: W'.

.::

*.*

1. -;

FIG. 13. Three hypotheses explaining the continuous variation between Myrceugenia glaucescens and M. ovata. A, phylogenetic continuum. B, parallel evolution. C, hybridization. (OV-M. ovata var. ovata; NA-M. ovata var. nannophvlla; GR-M. ovata var. gracilis; AT-M. ovata var. acutata; PI-M. pinifolia and other species; LT-M. glaucesens var. latior; GL-M. glaucescens var. glaucescens; CH-M. chrysocarpa; PL-M. planipes and other species).

tainty. Perhaps some future taxonomist with more material and additional data will study the group again. Key to Varieties of Myrceugenia glaucescens 1. Peduncles not subtended by bracts; leaves usually coriaceous, the midvein impressed proximally or not at all; apex and base usually not acuminate; calyx-lobes usually over 2 mm a. var. glaucescens. long; flowering mainly in December and January. 1. Some peduncles usually subtended by bracts; leaves usually submembranous, the midvein impressed for entire length or nearly so above; apex and base both acuminate; calyx-lobes b. var. latior. usually less than 2 mm long; flowering mainly in October.

20a. Myrceugenia glaucescens var. glaucescens

Figs. 14, 25, 3 lB.

Eugenia glaucescens Cambessedes, in Saint-Hilaire, Flora brasiliae meridionalis 2: 368, as to type, 1833. ?Eugenia araujoana Berg, in Mart. Fl. Bras. 14(1): 219. 1857. Type. Sellow s.n., "Habitat ad ripas fluminis Uruguay in Montevideo" (holotype B, apparently lost). Eugenia cambessedeana Berg, in Mart. Fl. Bras. 14(1): 230. 1857. Type. Sellow s.n., "Habitat ad ripas rivorum in Montevideo" (holotype, B, apparently lost; Field Museum neg. 36935 of probably isotype at P, photo at MICH). Eugenia bagensis Berg, in Mart. Fl. Bras. 14(1): 231. 1857. Type. Sellow s.n., type of E. bagensis [var.] a angustifolia Berg hereby designated as lectotype of E. bagensis Berg (lectotype, NY ex B). Eugenia bagensis [var.] a angustifolia Berg, in Mart. Fl. Bras. 14(1): 231. 1857. Type. Sellow s.n., "Habitat ad ripas amnis Rio Negro, prope Bag6, in Montevideo" (holotype, B, apparently lost; isotype NY ex B, hereby designated as lectotype). (Because the type of this taxon has been chosen as the lectotype of E. bagensis, the name would be replaced by E. bagensis var. bagensis).

Myrceugenia

Table V Mean Numerical Values for Characteristics Separating Myrceugenia glaucescens and M. ovata MEAN LOCULE NUMBER M. M. M. M.

glaucescens var. glaucescens ........................... glaucescens var. latior ................................ ovata var. acutata .................................... ovata var. gracilis .....................................

M. M. M. M.

glaucescens var. glaucescens ........................... glaucescens var. latior ................................. ovata var. acutata .................................... ovata var. gracilis ............................

M. M. M. M.

3.26 2.92 2.06 2.11

? + ? ?

0.43 0.70 0.09 0.18

MEAN STAMEN NUMBER

.........

183.3 136.0 71.7 64.5

? ? ? ?

24.6 35.6 58.9 11.5

7.73 6.15 5.10 4.08

+ + + +

0.74 1.00 0.97 0.97

MEAN OVULE NUMBER

?Eugeniabagensis [var.]/ avenia Berg, in Mart.Fl. Bras. 14(1):231. 1857.Type. Sellow s.n., "Habitatad ripas amnis Rio Negro, prope BagS, in Montevideo"(holotype, B, apparently lost). ?Eugenia bagensis [var.] y latifolia Berg, in Mart. Fl. Bras. 14(1): 231. 1857. Type. Sellow s.n.,

"Habitatad ripas amnis Rio Negro, prope Bage, in Montevideo"(holotype, B, apparently

lost).

?Eugeniapallida Berg, in Mart.Fl. Bras. 14(1):231. 1857.Type. Sellow s.n., "Habitatin Montevideo" (holotype, B, apparentlylost). ?Eugenia canelonensis Berg, in Mart. Fl. Bras. 14(1): 232. 1857. Type. Sellow s.n., "Habitat ad

ripas prope Canelonesin Montevideo"(holotype, B, apparentlylost).

?Eugenia elegans Berg, in Mart. Fl. Bras. 14(1): 232. 1857. Type. Sellow s.n., "Habitat in campis

ad Serpe in Montevideo,floret Decembri"(holotype, B, apparentlylost).

Eugenia ribeireana Berg, in Mart. Fl. Bras. 14(1): 307. 1857. Type. Sellow s.n., "Habitat in

Montevideo"(holotype, B, apparentlylost; probableisotype, P; Field Museumneg. 31612 of probableisotype at W, photo at MICH). Lumaglaucescens (Cambessedes)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 531. 1941. ?Lumaaraujoana(Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 532. 1941. Luma bagensis (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 532. 1941. ?Lumapallida (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 533. 1941. ?Lumacanelonensis (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 532. 1941. ?Luma elegans (Berg) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 533. 1941.

Luma angustior Burret, Notizbl. Bot. Gart. Berlin-Dahlem15: 530. 1941. Type. Bettfreund& Isolina Koster 782, "Argentinien,Buenos Aires, bei San Isidro" (holotype, B, apparently lost; isotype UC). ?Myceugenia pallida (Berg) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Mon-

tevideo 1(7):8. 1943.

?Myrceugenia glaucescens forma pallida (Berg) Legrand, Darwiniana 11(2): 345. 1957.

Myrceugeniaglaucescens forma debilis Legrandet Klein, Flora Ilustr. Catarinense[Mirt]:383. 1970. Type. Smith & Klein 13370, Santa Catarina,Cagador,MadeireiraRio Verde, 48 km ao oeste de Caqador,1000-1200m (holotype, HBR; isotypes, MICH, MVM, R, US). Myrceugeniaglaucescens formacatharinensisLegrandet Klein, FloraIlustr.Catarinense[Mirt]:

384. 1970. Type. Smith & Klein 13805, Santa Catarina, Campo Ere, 17 km ao oeste de Campo Ere, 900 m (holotype, HBR; isotypes, MICH, MVM, NY, R, SI, US). Myrceugenia grisea Legrand, in Legrand et Klein, Flora Ilustr. Catarinense [Mirt]: 387. 1970. Type. Smith & Klein 13122, Santa Catarina, Caibi, margem leste do Rio Iracema (holotype, MVM; isotypes, F, HBR, R, US).

Leaves 1.8-5.6 cm long, 0.6-1.8 cm wide, 2-8 times as long as wide, usually coriaceous, the midvein usually impressed only proximally or not at all; apex

Flora Neotropica

acute to rounded; base acute, cuneate or rounded;peduncles not subtendedby bracts; calyx-lobes 1.6-3 mm long, 1.6-3.5 mm wide. Type. Saint-Hilaire s.n., "Prope vicum S. Domingos Suriano in parte occidentali provinciae Cisplatinae"[Uruguay](holotype, P, n.v.; Field Museumneg. 36956 of probableholotype, photo MICH). Specimens examined. BRAZILParana:Mun. Curitiba,Uberabade Baixo (ca. 25?30'S,49?20'W), ca. 1000 m, 3 Nov 1977, Landrum2341 (MBM, MICH); Mun. Curitiba,rio Barigui(ca. 25?30'S, 49?20'W),ca. 900 m, 7 Nov 1977,Landrum2465,2466 (MBM,MICH).SantaCatarina:Mun.Cagador, 49 km west of Cacador, 1000-1200m, 18 Feb 1957, Smith & Klein 11366(US, R); Mun. Cacador, MadeireiraRio Verde, 48 km west of Cacador(ca. 26?45'S,51?22'W),1000-1200m, 2 Dec 1964, Smith & Klein 13351 (C, HBR, MO, MVM, R, US), Smith & Klein 13368 (HBR, MICH, MVM, R,

US); Mun. Chapec6, Fazenda Campo Sao Vicente, 24 km west of Campo Ere, 900-1000 m, 26-28 Dec 1956, Smith, Reitz & Sufridini9486 (HBR, MVM, R, US); Mun. Chapec6, 6 km west of Camp Ere, 900-1000 m, Smith & Klein 11525(HBR, MVM, R, US); Mun. Lajes, by Rio Bandeirinhas,23 km north of Lajes, 800-900 m, 4 Dec 1956, Smith & Klein 8232 (NY, R, US); Mun. Lajes, 4-6 km west of Capao Alto (ca. 27?50'S,50?30'W),ca. 1000m, 22 Nov 1977, Landrum2658b, 2659 (MBM, MICH); Mun. Lajes, ca. 18 km south of Lajes (ca. 27?50'S,50?20'W),ca. 1000 m, 22 Nov 1977, Landrum2681 (MBM, MICH);Mun. Xanxere, 5 km east of Fraxinaldos Guedes, 700-900 m, Smith & Reitz 9822 (HBR, MVM, R, US); Mun. Xanxere-Mun.Xaxim (ca. 26?55'S,52?29'W),700-800 m, Smith & Klein 13072(HBR, LP, MVM, NY, R, UC, US). Rio Grandedo Sul: Canela, 5 Feb 1948, A. Mattos & Labauriau s.n. (MICH, MVM, RB); Canela, Gramado, 26 Dec 1949, Rambo 45053 (H,

PACA); Caxias, Vila Oliva, 9 Jan 1946, Rambo 30805 (H, MVM, W), Rambo 30807 (H, MVM), 8 Feb 1955, Rambo56578, 56597 (MICH, MVM);Farroupilha,Santa Rita, 29 Jan 1949,Rambo40298 (H, PACA), 700 m, 7 Feb 1950,Rambo45706 (H, PACA);Vacaria,Passo do Socorro, 28 Dec 1951, Rambo51669 (MICH, MVM, PACA). URUGUAY. Artigas, 19 Jan 1942, RosengurttB 4011 (H); Dep. Canelones, rio Santa Lucia, estancia Paso Cuello, Feb 1946, Gallinalet al. PE 5580 (A, MVM, NY, PACA, SP); Cerro Largo, Sierrade Acegua, Jan 1919, Herters.n. (H); CerroLargo, Jan 1936,RosengurttA 799 (H), without date, RosengurttB 2743 (H, MVM);Colonia,Cascadadel ArroyoSan Luis, 10Nov 1946,Castellanos s.n. (W); Flores, Trinidad,100 m, Aug 1933, Herter 91377 (UC); Florida,EstanciaRinc6n de Santa Elena, 21 Apr 1943, Gallinalet al. PE 5188 (MO, MVM, NY); Florida,locality ?, Dec 1936,Rosengurtt B 799 (H, MVM); Minas, Salto del Penitente, Mar 1924, Herter s.n. (H); Maldonado,Cerro Pan de Azicar, Feb 1909, Arechavaleta s.n. (MVM, NY), ca. 400 m, Chebataroffs.n. (MICH, MVM), 6 Apr 1940, Legrand 2210 (H, MICH, MVM, NY), hacia la base, Legrand2211 (MICH, MVM), 300 m, 12 Mar 1946, Legrand4183, 4184 (H, MICH);Maldonado,Cerrodel Toro, 350 m, 4 Apr 1939, Legrand 1616 (H, MICH, MVM), 250 m, Legrand1613 (H, MICH, MVM, NY); Maldonado, Cerrode Animas, 8 Mar 1936, Legrand815 (H, MVM);Paysandu,rio Queguay,paso Andres Perez, Dec 1924, Schroeders.n. (H, MVM, NY); Rivera, Ataques, rio Tacuaremb6,Jan 1941, Legrand 2488 (H, MVM, NY); Rivera, Tranqueras,rio Tacuaremb6,Legrand 2485, 2486 (MICH, MVM), Osorio 32 (MICH, MVM, NY); Rocha, Arroyo Don Carlos, Legrand 1387 (MICH, MVM); Rocha, Arroyo Garz6n, Mar 1934, Legrand139 (H, MVM);Rocha, Rio Cebollati,frente a la Charqueada,20 Feb 1948, Castellanoss.n. (W); San Jose, Rinc6nde Arazati,24 Dec 1939,Legrand1708 (H, MVM); San Jose, rio Santa Lucia, Chebataroffs.n. (MICH, MVM); Soriano, Juan Jackson, estancia Santa Elena, Gallinal et al. 4504 (MICH), 24 Nov 1942, Gallinal et al. PE 4504 1/3 (H, MO,

MVM);Soriano, withoutlocality, 1 Jan 1941, RosengurttPE 4532 (H, MVM);Treintay Tres, Serra Yerbal, Quebradade los Cuervos, 9 Apr 1936, Legrand724 (H, MVM). PARAGUAY.Encarnaci6n,M bay Ca6, 19 Nov 1945,Bertoni2398 (MO, MVM). ARGENTINA. Misiones: Santa Ana, 5 Jan 1913, Rodriguez695 (H, MICH, MVM);Ap6stoles, Selva de Chirimay,29 Nov 1943,Burkart14301(MICH,SI); Candelaria,Garupa,9 Nov 1946,Bertoni 1999 (H, LIL, MVM). Corrientes:Santo Tome, EstablecimientoLas Marias,ruta nac. #14, 7 km S de gdor. Virasoro, 1 Dec 1970, Krapovickaset al. 16802 (SI). Entre Rios: Gualeguaychi, Cabrera 3972 (LP, MICH, MVM), 7 Jan 1932,Burkart4207 (SI); Dep. Uruguay,Lagrimade los Negros, 19 Dec 1941, Nicora 3229 (MICH, SI); Federaci6n, Martinez Crovetto & Leguizamou 5405 (MICH);

Dep. Federaci6n,rio Mocoreta,ruta 14, 15 Apr 1960,Burkart& Gamerro21840 (MICH,SI); Rinc6n del Mocoreta, 16Apr 1960,Burkart& Gamerro21844 (MICH, SI); orillasdel Mocoreta,9 May 1953, Nicora 6384 (MICH, SI), Nicora 6386 (SI); Delta, ParanaGuazu, 24 Nov 1932, Cabrera2510 (LP, MVM, NY); Delta, rio Brazo Largo, 7 Nov 1944,Jozami 256 (SI); Delta del Parana,Rio Ceibo, 25 Nov 1932,Burkart4905 (MVM, SI); Delta del Parana,Brazo Largoinferior,isla La Chilena, 1 Mar 1938,Burkart8947 (SI); Delta, ArroyoTuyupar6,1914,Scala 271 (NY), 272 (MVM, NY), 273, 274, 275 (LP), 276 (MVM). Buenos Aires: Las Conchas, Bonpland 1227 (RB); Atalaya FCS, Feb 1931, Burkart3675 (SI); Isla MartinGarcia, 19 Dec 1946,Palacios 13 (F, MVM);Delta del Parana,Burkart

Myrceugenia 70.0

-..:

*M/

FIG. 14. Distributionsof Mvrceugeniachrysocarpaand M. glaucescens.

4496 (MICH); La Plata, entre Rio Santiago y Palo Blanco, 2 Dec 1944, Rodriguez 578 (A); La Plata, Punta Lara, Dec 1938, Rodrigo 2219 (LP, NY), 4 Jan 1940, Dawson 928 (NY).

It is possible to deduce that Eugenia araujoana, E. pallida, E. canelonensis, and E. elegans are probably synonyms of M. glaucesens because of the descriptions, type localities and the fact that Burret, who probably saw the types, placed them in the genus Luma, which he understood to include Myrceugenia. Myrceugenia glaucescens var. glaucescens typically grows along streams from just south of Buenos Aires north to Parana. A map of its distribution is shown in Fig. 14. Towards the south the leaves are narrow and long whereas in the north they are shorter and broader (Fig. 25). In the south the plants are quite glabrous while in the north they tend to be somewhat pubescent. This clinal variation may be due in part to the hybridization that appears to take place with M. euosma in Santa Catarina and Paranai. This subject is discussed in the section on hybrids. Myrceugenia glaucescens var. glaucescens flowers mainly in December and January and the fruits mature shortly thereafter from February to April. This combination of summer flowering and fall fruiting is rare in the genus. This variety seems to be closely related to M. chrysocarpa of western South America and the two are compared at the end of Key M. Both are among the most cold-tolerant entities in the genus. 20b. Myrceugenia glaucescens var. latior (Burret) Landrum, Brittonia 32(3): 372. 1980. Fig. 14. Luma latior Burret, Notizbl. Bot. Gart. Berlin-Dahlem15: 530. 1941.

Flora Neotropica ...

.50....-

' S-..::.

' ^ 11.' 'S; tevideo 2(28): 1953.

' e :0-f *6::-^^^

,t,..-l',;|;

;

Levs -. c og,13cmwd, brnos mivenusalyimrssd the.:: acmnte FIG 15 aeusal Distributions ofcuiae;pdnle Mrceugenia rceugea

-. ..

-;s '; ;' d , h i . ;.CX .~~~~~~~~~~~~~~~~

* *:*\ f'-^i^-^

.-39tme sloga wd,summ o etielegh boe aexuual feseriatoramosa subtended and by :i?? calyxbat; cucullata M. scutellata

latior (Burret) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Mon-

branous, the midvein usually impressed for entire length above; apex usually acuminate; base usually acuminate; peduncles often subtended by bracts; calyxlobes 1.6-2.4 mm long, 1.2-2.2 mm wide. Type. Regnell III 570, "Minas Geraes, Caldas" (holotype, B, apparently lost; isotypes, C, GH, R). Specimens examined. BRAZIL. Minas Gerais: Pouso Alegre, a I km sul da encruzilhada Pouso Alegre-Itajuba-So Paulo, 30 Sep 1969, J. Mattos 15487 (S). Rio de Janeiro: Teres6polis, Serra dos

Myrceugenia

Orgaos, subida do Frade, 18 Jan 1883, Glaziou s.n. (R). Sao Paulo: Sio Paulo, 25 Sep 1918, Van Emelen 19 (SP, US); Cunha, 19 Oct 1939, Kiehl & Franco s.n. (SP). Parani: Mun. CerroAzul, Barra do Teixeira, 6 Oct 1960, Hatschbach 7311 (HB, HBR, MBM, MICH); Mun. Mangueirinha,Rio Igua9u, Porto Fanor, 20 Oct 1966, Hatschbach 15166 (MBM, MICH, MVM, US). Santa Catarina: Mun.Papanduva,Lajeadinho,750 m, 25 Oct 1962,Reitz & Klein13528(H, MVM,NY, UC, US); Mun. Papanduva,Picadas, ERF km 181, 750 m, 25 Oct 1962, Reitz & Klein 13535(H, MVM, US).

Myrceugenia glaucescens var. latior is known from relatively few collections

but as far as is known it has a more northerlydistributionthan M. glaucescens var. glaucescens. A map of its distributionis shown in Fig. 14. It is compared directly with M. ovata var. acutata in Key A. It flowers mainly in October and the only fruitingspecimen known was collected in September. 21. Myrceugeniascutellata Legrand, in Legrand et Klein, Flora Ilustr. Catarinense [Mirt]:436. 1970. Fig. 15. Tree ca. 6-10 m high; hairs reddish-brownto whitish, symmetricallydibrachiate, very small, inconspicuous; twigs glabrous to very sparsely pubescent, light grey to light reddish-brown;leaves elliptic, oblanceolate, or obovate, 2-8 cm long, 1-2(-3.5) cm wide, 2-3.5 times as long as wide, glabrous above, appearingglabrousbelow but covered with minute appressed hairs, these only visible with the compound microscope; apex acuminate, acute or rounded; base acute, cuneate, or acuminate;petiole channeled, 1-4 mm long, 0.5-1 mm wide, glabrousor nearly so, sometimes covered with thin, whitish, membranousscales, otherwise dark reddish-brown;midvein impressed for entire length or nearly so above, prominent below; lateral veins indistinct or up to ca. 10 pairs faintly visible; marginal veins about equallinglaterals in prominence;blade dryingreddish-brownto light green above, silvery light grey to reddish-tanbelow, coriaceous; peduncles 0.7-2 cm long, ca. 1 mm wide, glabrous, uniflorous,solitary or in pairs in the axils of leaves or at nodes where leaves have already fallen; bracteoles suborbicular, 1-2 mm long, somewhat broader than long, glabrous within and without; calyx-lobes broadly ovate, ca. 2-3 mm long, ca. 2-4 mm wide, ca. 0.7-1 times as long as wide, glabrouswithin and without; hypanthium ca. 1.5 mm long, broadly obconic, appearingglabrous when dry, but covered with a gelatinous matrix when wet, this matrix containing and covering hairs; disk sparsely puberulent, ca. 3-4 mm across; style sparsely pubescent to glabrous, ca. 5 mm long; stamens ca. 200-250, ca. 4-8 mm long; anthers ca. 0.50.8 mm long when dry; petals suborbicular,ca. 5-6 mm in diam.; ovary 3-locular in flowers seen; ovules 6-10 per locule; normalfruit unknown, probablypurple black, subglobose, ca. 1 cm in diam. Type. Reitz & Klein 11937, Santa Catarina,Lajes, Otacilio Costa (Encruzilhada), 900 m, 10 Jan 1962(holotype, MVM; isotype, HBR). Specimens examined. BRAZIL. Rio de Janeiro:Teresopolis, GranjaComari, 10 Nov 1964, A. Castellanos24541 (MVM). Parana:PaulaFreitas, 17 Nov 1972, Hatschbach30670 (NY); Mun. Gal. Carneiro,Basso da Galinha,19Nov 1972,Hatschbach30712 (NY). SantaCatarina:Mun.Anitapolis, Rio Branco, 600 m, 15 Dec 1972, Klein & Bresolin 10624(MVM).

This species seems to be similaron the one hand to Myrceugeniaglaucescens

and M. chrysocarpa and on the other to M. planipes, M. kleinii, M. reitzii, and

M. correifolia. It may represent a phylogenetic link between them or perhaps similarityto one of the groups is due to parallel evolution. When more material has been collected a better estimate of its position can be made. Myrceugeniascutellata is easily distinguishablefrom all the other species by its suborbicularbracteoles which are somewhat wider than long, and by its scar-

Flora Neotropica

city of hairs. It is known to flower in December and the fruits may matureshortly thereafter. A map of its distributionis shown in Fig. 15. 22. Myrceugeniacucullata Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 10 (name only) 1953;Darwiniana11: 347. 1957. Figs. 15, 31C. Shrub or tree up to 5 m high; hairs whitish to light reddish-brown,appressed, dibrachiate,more or less symmetric; twigs glabrousto densely pubescent when young, soon glabrescent; leaves glabrous to sparsely pubescent, elliptic to oblanceolate, 0.8-2 cm wide, 2.5-5 cm long, 2.5-3(-4.5) times as long as wide; apex acute to narrowlyacuminate; base cuneate to acuminate;petiole channeled, 24 mm long, 0.5-1 mm thick, sparsely to densely pubescent when young, glabrescent with age; midvein impressed proximallyor for entire length above, prominent below; lateral veins indistinctor up to about 20 pairsbarelyvisible; marginal veins equalling laterals in prominence; blades dull grey-green to yellow-green above, the same color but lighter beneath, membranousto subcoriaceous;peduncles uniflorous,flattened,0.5-4 mm long, 0.5-1 mm wide, densely pubescent, solitary or up to 3 in a row in the leaf axils; bracteoles lanceolate to ovate, 1.22.6 mm long, 0.7-1.6 mm wide, claspingthe hypanthiumor reflexed, membranous to subcoriaceous, densely to sparsely pubescent without and within; calyx-lobes ovate to ovate-triangular,concave, 1.5-3 mm long, 1.3-2.8 mm wide, 0.8-1.3 times as long as wide, subcoriaceous, densely to sparsely pubescent without, sparsely pubescent to glabrouswithin, the marginsoften apparentlyfusing at the apex to form a hornlike structurein 2 or more lobes, and/or the marginsof two lobes becoming quite thin and extending under the adjacentlobes; hypanthium obconic, 1.5-2.5 mm long, densely pubescent, sometimes extending about 1 mm beyond the ovary; disk 1.5-3.5 mm across, the staminalring pubescent to glabrous; style 4-6 mm long, sparsely pubescent; stamens 70-175(-230), 3-6 mm long; anthers 0.2-0.4 mm long when dry; petals suborbicular,concave, 2-2.5 mm in diam.; ovary bilocular or trilocular; ovules 4-8(-10) per locule; normal fruit unknown.

Type. Rambo 4319, Brazil, Rio Grande do Sul, Sao Francisco de Paula (holotype, MVM, isotype, PACA). Specimens examined. BRAZIL. Parana:Mun. Guaratuba,Rio Itarar6,31 Jan 1960, Hatschbach 6715 (HB, MBM, MICH,MVM);Mun. Piraquara,Mananciaisda Serra,950 m, 21 Feb 1968,Hatschbach 18633 (C, HB, MBM, MICH, MO, MVM, UC); Mun. QuatroBarras, Rancho Velho, 11 Feb 1964, Hatschbach 10968 (MVM). Santa Catarina:Mun. Lajes, 13 km east of Capao Alto, 900-1000 m, 12 Feb 1957, Smith & Klein 11326(HBR, MVM, R, US); Mun. Lajes, 6 km west of CapaoAlto, ca. 1000m, 22 Nov 1977, Landrum2666, 2671 (MICH, MVM);Nova Teutonia,27 Dec 1943, Plaumann 275 (RB); Mun. Vidal Ramos, Sabia, 750 m, 30 Dec 1957, Reitz & Klein 5942 (C, H, HBR, MVM, NY, US). Rio Grande do Sul: Taimbezinho,20 Feb 1953, Rambo 53979 (MICH, MVM, PACA); Sao Francisco de Paula, 7 Feb 1948, A. Mattos & Labauriau288 (MVM, RB); Mun. Sao Francisco de Paula, 5 km towards Porto Alegre from Sao Francisco de Paula, ca. 1000 m, 26 Dec 1977, Landrum2953, 2960 (MICH, PACA); Mun. Sao Francisco de Paula, Cascatinhanear Sao Franciscode Paula,ca. 1000m, 26 Dec 1977,Landrum2982 (MICH,PACA);Mun.Rolante,Fazenda Englertnear Santa Tereza, ca. 1000m, 28 Dec 1977, Landrum3004 (MICH, PACA).

Legrandtook up the name Myrceugeniaestrellensis (Berg) Legrandet Kausel for this species in the Flora Ilustrada Catarinense (1970), believing it to be an earlier name for his own M. cucullata. In the publicationin which Legrandand Kausel transferred Eugenia estrellensis to Myrceugenia (Legrand, 1953) and in

Legrand's treatmentof the eastern species of Myrceugenia (1957), Legrand expressed the opinion that M. estrellensis was distinct from M. cucullata. Unfortunatelyno confirmabletype specimenhas been foundfor E. estrellensis although

Myrceugenia

a possible isotype has been located at P. That specimen belongs to M. myrcioides and Berg's description also suggests M. myrcioides. FurthermoreI know of no collections of M. cucullata north of Parana, so it does not seem likely that it reaches the type locality of E. estrellensis at Serra da Estrella in Rio de Janeiro. ThereforeI have not followed Legrandin using the name M. estrellensis for this species.

Luma mesomischa Burret(1941a)may be an earliername for M. cucullata but unfortunatelyno type has been found and the descriptionis too ambiguousto be sure what species Burretwas describing. Myrceugenia cucullata is a puzzling species, being somewhat similar to M. ovata var. acutata, M. seriatoramosa, and M. myrcioides var. acrophylla. What

its true phylogenetic affinities are I do not know. With the first two it has been compared directly in the key. The differences between it and the last are summarizedbelow. M. cucullata

M. myrcioides var. acrophylla

Leaves 2.5-5 cm long Calyx-lobes 1.5-3 mm long Twigs glabrous to densely pubescent when young, soon glabrescent Stamens 70-175(-230).

Leaves 4-16 cm long Calyx-lobes(2.5-)4-6.5 mm long Twigs densely pubescent, not soon glabrescent Stamens250 to more than 500

Myrceugenia cucullata is rather common in the Araucaria forests of southern

Santa Catarinaand in Rio Grandedo Sul. A map of its distributionis shown in Fig. 15. It flowers from December to Febraury, and although undiseased fruits are unknown, those attacked by a fungus may persist up to a year on the plant. 23. Myrceugeniaseriatoramosa(Kiaerskou)Legrandet Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 5. 1953. Figs. 15, 31D. Eugenia seriato-ramosaKiaerskou,Enum. Myrt. bras.: 170. 1893. Luma seriato-ramosa(Kiaerskou)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 528. 1941.

Tree up to 5 m high; hairs reddish-brown, symmetrically dibrachiate, appressed; twigs moderatelypubescent to glabrous, the young bark light reddishbrown, becoming whitish-greywith age; leaves sparsely pubescent to glabrous below, completely glabrous or sparsely pubescent along the midvein above, elliptic to ovate or obovate, 2.8-5.7 cm long, 1-2.7 cm wide, 1.9-3 times as long as wide; apex acute or acuminate, the tip usually blunt; base acuminateto cuneate; petiole channeled, 3-7 mm long, ca. 1 mm thick, sparsely pubescent to glabrous; midvein impressed for entire length or only proximallyabove, prominent below; lateral veins usually indistinct, up to 16 pairs, sometimes faintly visible; marginal veins equallinglaterals in prominence; blades light grey-green or becoming dark reddish-brownupon drying, often lustrous above, dull yellowgreen or lighter brown below, coriaceous; peduncles uniflorous,flattened, 0.4-1 cm long, ca. 0.5-1 mm wide, densely pubescent, solitary or 2-3 in a row in the leaf axils; bracteoles ovate to lanceolate, 1.7-4.3 mm long, 1-2 mm wide, 1.32.2 times as long as wide, coriaceous to membranous,densely to sparsely pubescent without, sparsely pubescent to glabrous within, loosely clasping the hypanthium;calyx-lobes ovate to suborbicular,1.7-4 mm long, 1.2-3 mm wide, 11.5 times as long as wide, concave, coriaceousto membranous,densely to sparsely pubescent without, sparsely pubescent to glabrouswithin; hypanthiumobconic, densely pubescent, 1.5-3 mm long; disk 2-4 mm across, sparsely pubescent; style 3-8 mm long, sparsely pubescent; stamens 80-240, 2-8 mm long; anthers

Flora Neotropica

0.3-0.6 mm long when dry; petals concave, suborbicular, 3-5 mm in diam.; ovary 2-3-locular; ovules 5-14 per locule; fruit unknown. Type. Glaziou 17006, "Rio de Janeiro" (holotype, C, n.v.; isotype, US; Field Museum neg. 20977 of holotype, photo at MICH). Specimensexamined.BRAZIL.Rio de Janeiro:Campodas Antas, 2200m, 20 Apr 1949,Alt. Barb. 119 (RB). Parana:Carvalho,7 Nov 1911,Dusen 13324(MICH,NY); Mun. Piraquara,Rio do Corvo, 1 May 1949, Hatschbach 1393 (MBM, MICH, MVM, PACA); Mun. Piraquara,MorroAnhangava, 8 Apr 1951,Hatschbach2226 (HBR, MBM, MICH, MVM);Mun. QuatroBarras,RanchoVelho, 23 Feb 1964, Hatschbach 10991 (MBM, MVM).

Myrceugenia seriatoramosa might be confused with and is perhaps most closely related to M. alpigena var. rufa, which differs in having the lower leaf surfaces, the young twigs and the calyx-lobes densely covered with hairs. The phenology of Myrceugenia seriatoramosa is unclear because so few specimens have been collected, but it appears to flower in April and May. Myrceugenia seriatoramosa grows from Rio de Janeiro to Parana, apparently at fairly high elevations near the coast. A map of its distribution is shown in Fig. 15. 24. Myrceugenia colchaguensis (R. A. Philippi) Navas, Bol. Mus. Nac. Hist. Nat. Chile 29: 230. 1970. Figs. 17, 32A. Eugenia colchaguensis R. A. Philippi, Linnaea 33: 72. 1864. Mvrceugenia malvillana Kausel, Lilloa 17: 51. 1949. Type. Kausel s.n., "Valle de Malvilla (Chile,

prov. de Santiago),"(holotype, H).

Shrub 2-4 m high; hairs golden-brown to yellowish-brown, more or less symmetrically dibrachiate, twisted; twigs densely lanate when young, glabrescent with age, strongly flattened at the nodes; leaves densely lanate below when young, less densely so to glabrous above when young, glabrescent on both surfaces with age, elliptic to obovate, 0.7-2(-3) cm long, 0.4-1.2(-1.5) cm wide, 1.42 times as long as wide; apex obtuse to rounded; base rounded to acute; petiole 1-2 mm long, ca. 1 mm thick, densely lanate when young, glabrescent with age, slightly channeled; midvein recessed proximally or not at all above, prominent proximally, indistinct distally below; lateral and marginal veins indistinct; blades light yellow-green to grey-green, lustrous or dull above, lighter beneath, coriaceous to subcoriaceous; peduncles uniflorous, flattened, 0.8-1.7 cm long, ca. 1 mm wide, densely lanate, solitary or two together in the leaf axils; bracteoles lanceolate, 1.5-2.5 mm long, 0.7-1 mm wide, 2-2.8 times as long as wide, densely lanate within and without, subcoriaceous, clasping the hypanthium; calyx-lobes ovate to broadly triangular, 2-3 mm long, 2-3.5 mm wide, 0.7-1.2(-1.4) times as long as wide, densely lanate distally to glabrous proximally within, densely lanate without, coriaceous, usually somewhat concave; hypanthium densely lanate, 1.22 mm long; disk 2-3 mm across, sparsely covered with hairs; style 5-9 mm long, sparsely pubescent to glabrous; stamens ca. 180-240, 5-8 mm long; anthers ca. 0.5 mm long when dry; petals concave, more or less orbicular, ca. 3-5 mm in diam.; ovary usually 3-locular; ovules 6-12 per locule; mature fruit unknown. Type. Landbeck s.n., "In provincia Colchagua" (holotype, SGO-064324; probable isotype, MICH ex G; Field Museum neg. 23529 of probable isotype at P, photo at MICH). Specimensexamined.CHILE. Santiago:ca. 2 km west of Malvilla,ca. 10 km east of San Antonio

(ca. 33?35'S, 71?31'W), ca. 100 m, 2 Apr 1978, Landrum 3454, 3455, 3456, 3457, 3458, 3459, 3460 (MICH, SGO). Colchagua: La Serena, La Rufina, 5 Jan 1951, Ricardi 10112 (CONC). Talca: Camino de Talca a Curepto, km 65, 400 m, 12 Jan 1964, Marticorena & Matthei 495 (CONC). Maule: Con-

Myrceugenia

stituci6n, Feb 1891, Reiche s.n. (SGO). Malleco: Fundo Sto. Domingo, Inspector Fernmndez, 10 Jan 1955, Pfister 18543 (CONC).

Myrceugenia colchaguensis has been collected less than any other western species of Myrceugenia. Although it is actually quite distinct, it has been confused with other species such as M. correifolia and M. rufa. Where I collected it, near Malvilla, it was growing with M. obtusa and was quite similar to that species in general appearance. Myrceugeniacolchaguensis can be distinguished from the other western species by the combinationof calyx-lobes that are densely lanate without; yellowish, symmetricallydibrachiate,twisted hairs; small leaves; and uniflorouspeduncles. Its closest relative is probably M. alpigena of Brazil. M. colchaguensis is comparedto that species in the discussion of M. alpigena var. rufa.

Myrceugenia colchaguensis flowers in the summer months of January and February. Mature fruits are unknown. It is known from only five localities in Chile. A map of its distributionis shown in Fig. 17. 25. Myrceugeniafranciscensis(Berg) Landrum,Brittonia32(3): 372. 1980. Figs. 18, 32B. Eugenia franciscensis Berg, in Mart., Fl. Bras. 14(1): 233. 1857. Luma mucronata Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 533. 1941. Type. "Brasilien: Ilha do Mar (Herb. Sao Paulo n. 27019 cor F. C. Hoehne)" (holotype, SP). Myrceugenia mucronata (Burret) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 10. 1953. Mvrceugenia bracteosa var. franciscensis (Berg) Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 7. 1953. Mvrceugenia longipedunculata Mattos et Legrand, Loefgrenia 67: 18. 1975. Type. Hatschbach 18950, Parana, Mun. Jaguariaiva, Barra do rio das Mortes, 24 Feb 1968 (holotype, MBM, seen at MVM; isotype, UC).

Shrub ca. 1.5 m high; hairs reddish-brownto yellowish-brown,symmetrically dibrachiate, somewhat spreading; twigs densely pubescent when young, losing most or all pubescence before the first bark falls, the bark greenish-grey,tan to light tan when young, remainingsmooth; leaves more or less elliptic, (2-)4-6.5 cm long, 1.3-3.5 cm wide, ca. 1.2-2.5 times as long as wide, densely pubescent on both surfaces when young, glabrescent or nearly so at maturity;apex acute to rounded, sometimes mucronate;base acute to acuminate;petiole channeled, densely pubescent at first, glabrescent with age, 3-5 mm long, 1-1.5 mm thick, light yellow to grey or reddish-brownwhen dry; midvein impressed only slightly proximally or for entire length above, prominentbelow; lateral veins indistinct to moderately prominent, up to ca. 20 pairs visible; marginal veins equalling lateralsin prominence;blades light grey-greento yellow-greenor dryingreddishbrown above, somewhat lighter below, subcoriaceous to coriaceous; peduncles flattened,densely pubescent (glabrescentwith age?), (1-)2-4.5 cm long, ca. 1 mm wide; bracteoles ovate to lanceolate, ca. 3-4 mm long, ca. 2 mm wide, densely pubescent without, densely pubescent distally, less so proximallywithin; calyxlobes ovate, apparentlyone pair or all four blunt, ca. 2-4 mm long and wide, densely pubescent within and without; hypanthiumobconic, densely pubescent, ca. 2 mm long; petals apparentlyglabrous; ovary 3-locular; ovules ca. 6-7 per locule. Type. Martius s.n., "Habitat ad flumen S. Francisci: M.," "(v. in hb. Mart. et Sonder.)" (isotype at M hereby designated as lectotype; paratype at MEL, n.v.). Additional specimen examined. BRAZIL. Sao Paulo: rodovia Itarare-Bom Sucesso, a 15 km de Itarare, 20 Oct 1967, J. Mattos 13993 (SP).

Flora Neotropica

This species is known from only four collections, none of which is complete. Unfortunately, it has no outstandingmarkercharacteristics. It is very close to Myrceugenia alpigena and M. euosma, and in certain characteristics seems to

bridgethe gap between them. As more materialaccumulatesit may become clear that M. franciscensis should be reduced to a variety of one or the other of these

species, but for the present I believe it preferableto retainit as a separatespecies. It is comparedwith M. euosma and M. alpigena in the table below. M. franciscensis

M. euosma

Leaves glabrescent or nearly so at maturity, Leaves densely pubescent beneath at matu(2-)4-6.5 cm long, 1.3-3.5 cm wide. 1.2rity, 0.8-3(-4.7) cm long, 0.2-1.2(-1.5) 2.5 times as long as wide cm wide, (1.7-)2-6 times as long as wide Peduncles(1-)2-4.5 cm long Peduncles0.4-1.2 cm long Hairs on twigs all dibrachiate Hairs on twigs a mixtureof simple and dibrachiate M. franciscensis

Leaves glabrescent or nearly so at maturity, (2-)4-6.5 cm long, 1.3-3.5 cm wide, 1.22.5 times as long as wide Peduncles(1-)2-4.5 cm long Hypanthiumaboutas longas or shorterthanthe calyx-lobes Bracteolesca. 1.5-2 times as long as wide

M. alpigena var. alpigena

Leaves densely pubescentbeneathto glabrescent at maturity,1-4(-4.6) cm long, 0.31.5 cm wide, 2-3.7 times as long as wide Peduncles0.6-2.2 cm long Hypanthiumusually longer than the calyxlobes Bracteolesca. 2-4 times as long as wide

M. franciscensis

M. alpigena var. rufa

Leaves glabrescentor nearly so at maturity, Leaves densely pubescent beneath at maturimore or less elliptic ty, usually obovate Pedunclessolitaryor in pairsin the leaf axils, Pedunclesusually 2-4 togetherin the leaf axils, 0.3-1 cm long (1-)2-4.5 cm long

The phenology of M. franciscensis is still unclear but it probably flowers in November or December. A map of its distributionis shown in Fig. 18. 26. Myrceugeniaalpigena (A. P. de Candolle) Landrum, Brittonia 32(3): 372. 1980. Eugenia alpigena A. P. de Candolle,Prodr.3: 265. 1828.

Tree or shrub 2-6 m high; hairs reddish-brownto greyish or yellowish-brown, symmetricallydibrachiate,the arms straightand appressed or less often curled and twisted together; twigs densely strigose to tomentose when young, glabrescent with age, some hairs usually persistinguntil the first bark exfoliates; leaves elliptic, narrowly elliptic, oblanceolate, obovate, lanceolate or ovate, 1-5.5(-9) cm long, 0.3-2(-2.5) cm wide, 2-3.7(-5) times as long as wide, usually densely covered with hairs below when young, less often sparsely so, moderately to densely covered with hairs above when young, glabrescenton both surfaces with age; apex acuminate, acute, obtuse, or rounded; base cuneate, acute, or acuminate;petiole channeled, 2-6 mm long, 0.5-1.5 mm thick, densely covered with hairs when young, glabrescent with age; midvein impressed for entire length, only proximally, or not at all above, prominentbelow; lateral veins indistinctor up to ca. 10 pairs faintly visible; marginalveins equallinglateralsin prominence; blades reddish-brown, yellowish-brown, grey-green to yellow-green above, somewhat lighter below, coriaceous (rarely subcoriaceous), the upper surface usually dull; peduncles uniflorous(rarely with a three flowered dichasium),flattened or sulcate, 3-22 mm long, 0.5-1 mm wide, usually densely covered with

Myrceugenia ..

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FIG. 16. Scatter diagramof Myrceugeniaalpigena. Calyx-lobe ratio equals calyx-lobe length/ calyx-lobe width. Bracteole ratio equals bracteole length/bracteolewidth. All numericalvalues are averages of two to several observations.

hairs, solitary or up to 4(-5) in the leaf axils; bracteoles orbicular, ovate or lanceolate, 1.4-4 mm long, 0.8-2.9 mm wide, 0.9-4 times as long as wide, submembranous to coriaceous, densely covered with hairs without, sparsely to densely covered within, clasping the hypanthium; calyx-lobes orbicular, ovate to triangular, 1.6-3.2 mm long, 1-3.3 mm wide, 0.8-1.7 times as long as wide, more or less concave, submembranous to coriaceous, densely covered with hairs with-

Flora Neotropica

out, sparsely to densely covered with hairs within; hypanthiumobconic, 1.5-3 mm long, densely covered with hairs; disk 2-3 mm across, moderatelypubescent; style 5-7 mm long, sparsely pubescent to glabrous; stamens 80-180, 4-8 mm long; anthers 0.3-0.5 mm long when dry; petals glabrous or nearly so, suborbicular, 4-5 mm in diam.; ovary 2-4-locular; ovules 4-12 per locule; fruit not well known, ca. 5-6 mm in diam., apparentlyoften attacked by a fungus, this formingnumeroussmall black spheres on the surface. As defined here, Myrceugeniaalpigena is an inclusive species with three well marked varieties between which there exist some transitionalspecimens, especially in the area of Itatiaia. The scatter diagramin Fig. 16 summarizesthe morphological variationin the species. Two collections made by Dusen (528 and 2027) from Itatiaia have been provisionally placed in Myrceugeniaalpigena var. rufa. They show characteristics of all three varieties but seem to approachthis one most closely. It is possible that they represent a new entity but the differences are too subtle to make a judgment with so few specimens. The Myrceugenia alpigena complex, which includes M. bracteosa, M. franciscensis, M. euosma, M. seriatoramosa and M. colchaguensis, is a difficult one. Myrceugenia alpigena is compared to M. franciscensis in the discussion of the

latter species. But otherwise each variety of M. alpigena is comparedwith those members of the complex with which it might most easily be confused in the discussion of this species. Key to Varieties of M. alpigena 1. Leaves mainly over 5 cm long, 3-5 times as long as wide, sparsely pubescentto glabrous c. var. longifolia. beneath;Goias near Brasilia. 1. Leaves mainly less than 5 cm long, mostly 2-3 times as long as wide, usually densely pubescent beneath at least when young; Minas Gerais and Espirito Santo south to Rio Grandedo Sul. 2. Calyx-lobesacute, about as long as the hypanthium;bracteoles mainly 2 times as long a. var. alpigena. as wide. 2. Calyx-lobesbluntto rounded,1.3-1.7 times as long as the hypanthium;bracteolesmainly b. var. rufa. less than 1.7 times as long as wide.

26a. Myrceugenia alpigena var. alpigena Eugenia alpigena A. P. de Candolle,Prodr.3: 265, as to type, 1828. Psidium itatiaiae Wawra,ItineraPrincipumS. Coburgi1: 20. 1883.Type. Wawra I 483, "Brasilien. Granitzinkende Itatiaia"(holotype, W, n.v.; Field Museumneg. 31424of holotype, photo at MICH). Luma alpigena (A. P. de Candolle)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 535. 1941. Mvrceugeniabracteosa var. alpigena (A. P. de Candolle)Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo2(28):7. 1953. MyrceugenialeptorhynchaLegrand,in Legrandet Klein, Flora Ilustr. Catar. [Mirt]:392. 1970. Type. Reitz & Klein 10496, Santa Catarina,Garuva,Morrodo CampoAlegre, 1200m, 21 Dec 1960(holotype, MVM;isotype, HBR). Mvrceugenia bracteosa var. guaratubensis Mattos, Loefgrenia 65: 6. 1975. Type. Hatschbach

6684, Parana,Guaratuba,Serrade Aracatuba,1350m, 31 Jan 1960(holotype, MBM). Myrceugeniacatharinae Legrand, Flora Ilustr. Catar. [Mirt-Supl.]:30. 1977. Type. Klein & Bresolin 10089, Santa Catarina,Morrodo Cambirela,700 m, 23 Feb 1972(holotype, MVM; isotype, HBR).

Leaves 1-4(-4.6) cm long, 0.3-1.5 cm wide, 2-3.7 times as long as wide, mainly elliptic, usually densely covered with hairsbeneath;peduncles solitaryor in pairs in the leaf axils, 0.6-2.2 cm long; bracteoles lanceolate, 2-4 mm long, 0.8-1.8 mm wide, 2-4 times as long as wide; calyx-lobes acute, triangularto triangular-

Myrceugenia

ovate, 1.6-3.2 mm long, 1-3 mm wide, 0.9-1.7 times as long as wide; hypanthium 1.5-3 mm long.

Type. Martiuss.n., "in montibusBrasiliaeprov. Minarum,""v. s. sine fr. in h. Mart." (holotype, M; Field Museumneg. 19915of holotype, photo at MICH). Specimensexamined.BRAZIL.EspiritoSanto:Pico da Bandeira,Serrado Capara6,ca. 9200ft, 3 Mar 1959,Irwin2778 (MICH,R). MinasGerais:Serrado Capara6,perto do Rio Claro, 29 Nov 1960, Vidal 97 (R); Serra do Capara6,Macieiras, 15 Nov 1960, Strang 225 (GUA, MVM, RB). Rio de Janeiro:Itatiaia,AgulhasNegras, Feb 1938, CamposPorto 3423, 3424 (MVM, RB), 2800 m, 27 May 1935, Brade 14609 (RB), 2600 m, May 1950, Brade 20333 (MVM, RB); Itatiaia, Cume das Agulhas Negras, 21 Oct 1922, Kuhlmanns.n. (RB); Itatiaia,planalto, 24 Feb 1943, Pereira 20 B (HB, RB); Itatiaia, planalto, subida das Agulhas Negras, 2400-2500 m, 6 Feb 1969, Sucre 4655 (MICH, RB); Itatiaia, estrada nova, km 8, 25 Mar 1942, Brade 17264 (RB); Itatiaia,road to AgulhasNegras, km 8, ca. 2300 m, 18 Oct 1977, Landrum2110, 2112 (MICH, RB); Itatiaia, km 21, Jan 1939, Lanstyak 252 (RB, US); Parque Nacional de Itatiaia, 13 Feb 1960, Barth F. 190 (US); Campos de Itatiaia, withoutdate, Glaziou6544 (C, F, RB); Teres6polis, Serrados Orgaos, Pedrado Sino, 2150 m, Feb 1952, Vidal 11-705(R); Haut dos Orgaos, Mar 1884, Glaziou 14834 (C, US), 21 Jan 1887, Glaziou 16050(C, F, US).

Myrceugeniaeuosma might be confused with this variety but it has a mixture of simple and dibrachiatehairs and the bracteoles are mainly less than twice as long as wide, while in M. alpigena var. alpigena the hairs are all symmetrically dibrachiateand the bracteoles are usually 2-4 times as long as wide. In the area where their ranges come nearest to each other in Paranaand Santa Catarina,M. euosma grows towardsthe interiorof the planaltoin the Araucariaforests whereas M. alpigena var. alpigena seems to be restricted to the high elevations and misty forests of the eastern edge of the planalto. If plotted on the scatter diagram in Fig. 16, M. euosma would appearin the lower right hand corner. Myrceugeniaalpigena var. alpigena has mainly been found in Itatiaia and in Serra dos Orga6s in Rio de Janeiroat elevations above 2000 m but has also been collected on Pico da Bandeira in Espirito Santo at a similar elevation, and in Guaratuba,Parana and Garuva, Santa Catarinaat 1200-1350 m. A map of its distributionis shown in Fig. 17. It clearly needs a cool moist habitat. It flowers from Januaryto May and its fruits probablymaturein October and November. 26b. Myrceugeniaalpigenavar. rufa (Berg) Landrum,Brittonia32(3): 372. 1980. Figs. 17, 32C. Eugeniafuliginea [var.] 3 rufa Berg, in Mart., Fl. Bras. 14(1):571. 1859. ?Eugeniafuliginea Berg, in Mart., Fl. Bras. 14(1): 233. 1857. Type. Sellow s.n., "Habitat in Brasiliameridionali"(holotype, B, apparentlylost). Eugenia seriato-pedunculata Kiaerskou, Enum. Myrt. bras.: 170. 1893. Type. Glaziou 14821,

locality accordingto Glaziou(1905-1913),MinasGerais, "Itacolumy,pres OuroPreto" (holotype, C; Field Museumneg. 20976, photo of holotype at MICH).

Eugenia sticheromischa Kiaerskou, Enum. Myrt. bras.: 172. 1893. Type. Glaziou 14833, locality

accordingto Glaziou(1905-1913),MinasGerais, "Agua Limpa,h Gandarela"(holotype, C; Field Museumneg. 20979of holotype, photo at MICH). Lumaseriato-pedunculata(Kiaerskou)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 527. 1941. Luma sticheromischa(Kiaerskou)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 526. 1941. Myrceugenia bracteosa var. seriato-pedunculata (Kiaerskou) Legrand, Comun. Bot. Mus. Hist.

Nat. Montevideo2(28):9. 1953. Myrceugeniabracteosa var. australis Legrand,Comun.Bot. Mus. Hist. Nat. Montevideo2(28): 7 (name only). 1953;Darwiniana11(2):335 (description).1957;in Legrandet Klein, Flora Ilustr. Catar. [Mirt]:351 (type designated). 1970.Type. Rambo 56280, Rio Grandedo Sul, Sao Francisco de Paula, Fazenda Englert, 2 Jan 1955 (holotype, MVM; isotypes, MICH, PACA).

Leaves 1.5-5.5 cm long, 0.6-2 cm wide, 2-3 times as long as wide, mainly obovate, usually densely covered with appressed hairs beneath; peduncles usu-

Flora Neotropica

ally 2-4 in a row in the leaf axils, 0.3-1 cm long; bracteoles ovate to orbicular, 1.4-2.4 mm long, 1-2.8 mm wide, 0.9-1.8 times as long as wide; calyx-lobes blunt to rounded, ovate to orbicular, 1.6-2.7 mm long, 1.4-3.3 mm wide, 0.8-1.2 times as long as wide; hypanthium1.5-2 mm long. Type. Riedel s.n., "Habitat in fruticetis ad colles editos prope Ouro Preto prov. Minarum"(holotype, LE; probableisotype, P). Specimensexamined. BRAZIL. MinasGerais:Serrado Cip6, Serrado Espinhaco,ca. 1125m, 18 Feb 1972, Anderson, Stieber & Kirkbride36240 (MICH);Pico de Itacolomi, ca. 2 km S of Ouro Preto, Serra do Espinhago, 1600 m, 30 Jan 1971, Irwin, Harley & Onishi 29383, 29386 (MICH); Itacolumy,pres OuroPreto, Glaziou 19363a(C); pres Barbacena,Dec-Jan, Glaziou 16066(C, RB); Serrade Sao Jose d'El Rei, Oct-Nov, Glaziou14271(C, MVM);Serrade OuroPreto, Feb 1892, Ule 2493 (R). Rio de Janeiro:Tijuca, 7 Dec 1928,Pessoal do Horto Florestal s.n. (MICH, RB). Parana: Mun. Tijucas do Sul, Rincao, 13 Jan 1963, Hatschbach 9696 (HB, MBM, MICH, MVM); Mun. Palmeira,Serradas Almas, 12Jan 1966,Hatschbach13483(MBM, MICH, MVM, US), Hatschbach 13482(F); Mun. Sao Jose dos Pinhais,Contenda,20 Dec 1967,Hatschbach18158(HB, MBM,MICH, MO, MVM,NY, UC); Mun. Sao Jose do Pinhais,Rod. Br. 277, Rio Itaqui,28 Dec 1972,Hatschbach 31065 (C, HB, MBM, NY, UC). Santa Catarina:Mun. Mafra,ca. 10 km west of Rio Negrinhoat bridgecrossing Rio Preto (ca. 26?15'S,49?37'W),ca. 900 m, 24 Nov 1977, Landrum2722, 2723, 2730 (MBM, MICH);Mun. CampoAlegre, Morrodo Iquererim,1300m, 5 Feb 1958, Reitz & Klein 6435 (H, HBR, MVM, US), 9 Jan 1958,Reitz & Klein6024 (HBR, MVM, US), 10Jan 1958,Reitz & Klein 6160 (H, HBR, MVM, US); Mun. CampoAlegre, lower fazenda of Ernesto Scheide, ca. 900 m, 1 Feb 1957,Smith & Klein 10563(MICH, MVM, NY, R, RB, US); Mun. Matos Costa, south of Matos Costa on the road to Calmon(22 km), 900-1100 m, 20 Dec 1956, Smith & Reitz 8921 (HBR, MVM, R, US); Mun. Sao Joaquim,8 km south of Sio Joaquim(ca. 28?20'S,49?56'W),1300m, 5 Jan 1965, Smith & Reitz 14281 (HBR, MICH, MVM, R, US); Mun. Sao Joaquim, Tres Pedrinhas, 12 km southwest of Sao Joaquim(ca. 28?22'S,49?57'W),1200-1300m, 6 Jan 1965, Smith & Klein 14344 (HBR, MICH, MVM); Mun. Bom Jardim,CurralFalso, 1500m, 19 Feb 1959, Reitz & Klein 8393 (HBR, MVM, NY, UC, US), 19 Mar 1959,Reitz & Klein8675 (H, HBR, MVM).Rio Grandedo Sul: Mun. Sao Francisco de Paula, Cascatihnanear Sao Francisco de Paula, ca. 1000 m, 26 Dec 1977, Landrum2962, 2961 (MICH,PACA);Taimbesinho,p. Sao Franciscode Paula, 20 Feb 1953,Rambo 54062, 54094 (MICH).

The name Myrceugenia bracteosa has been applied to this variety in the numerous works of Legrand. As definedhere, M. bracteosa is a distinct but rather similar species which can be distinguishedfrom M. alpigena var. rufa through its hairs which are a mixture of simple and dibrachiate, not all symmetrically dibrachiate;its calyx-lobes, at least two of which are usually strongly acute, not blunt to rounded; and its leaves which are mainly elliptic, often with the veins somewhat prominent,not obovate with indistinctveins as they usually are in M.

alpigena var. rufa. Myrceugenia seriatoramosa is also quite similar to M. alpigena var. rufa but

differs in having the lower surfaces of leaves, the twigs and sometimes the calyxlobes only sparsely covered with hairs. In M. seriatoramosa the leaf apex is usually bluntlyor sharplyacuminate,not roundedas it normallyis in M. alpigena

var. rufa. Myrceugenia colchaguensis, the only western member of the M. alpigena com-

plex, might also be confused with M. alpigena var. rufa if one did not know the origin of a specimen. It differs in that its leaves are only 0.7-2(-3) cm long and its bracteoles are 2-2.8 times as long as wide. It differs from M. alpigena var. alpigena in its calyx-lobes being 0.7-1.2(-1.4) times as long as wide and the hypanthiumbeing 1.2-2 mm long. If plotted on the scatter diagramin Fig. 16 it would mainly be located to the left of M. alpigena var. alpigena and above M.

alpigena var. rufa.

Myrceugenia alpigena var. rufa ranges as far north as Minas Gerais but apparently becomes more common towards the south until at its southern limit in Rio Grande do Sul it is one of the main constituents of the Araucaria forests.

Myrceugenia

There it may play a pioneer role in the expansion of the forests because it commonly grows around the edges. It flowers from December to March. Normal fruits are unknownto me but those attackedby fungi seem to persist on the plants for at least a year. A map of the distributionof M. alpigena var. rufa is shown in Fig. 17. 26c. Myrceugeniaalpigenavar. longifolia(Burret)Landrum,Brittonia32(3): 372. 1980. Fig. 17. Luma longifolia Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 532. 1941. Myrceugenialongifolia (Burret)Legrandet Kausel, in Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 5. 1953.

Leaves 4.5-9 cm long, 1.3-2.5 cm wide, 3.5-5 times as long as wide, oblanceolate, sparsely pubescent to glabrous beneath; peduncles usually 1 or 2 in a row in the leaf axils, 0.5-1.5 cm long; bracteoles ovate, 2.2-3.2 mm long, 1.32.9 mm wide, 1-1.7 times as long as wide; calyx-lobes rounded, ovate to oblong, 2.6-3.2 mm long, 2.2-2.8 mm wide, 0.9-1.4 times as long as wide; hypanthium ca. 2-2.5 mm long. Type. Glaziou21147, "Brasilien:Goyaz, zwischen Rio Torto und Bananal,im campo, Strauch mit weissen Bluten" (isotype, C). Additionalspecimen examined. BRAZIL. Goias: Formosa, margemda Lagoa Feia, 17 Oct 1964, Heringer 9961 (MVM).

This variety seems to be endemic to the vicinity of Brasilia. A map of its distributionis shown in Fig. 17. Perhaps an ancestor migratedto this northern outpost of the genus Myrceugenia during the Pleistocene. At present the area aroundBrasilia seems to be an island of relatively cool temperaturessurrounded by an area of warmerclimate. 27. Myrceugeniabracteosa(A. P. de Candolle) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 6. 1953. Fig. 18. Eugenia bracteosa A. P. de Candolle,Prodr.3: 276. 1828. Eugenia montana Cambessedesin Saint-Hilaire,Flora Brasiliaemeridionalis2: 365. 1833.Type. Saint-Hilaires.n., "In dumetisvulgoCarrascossummimontisSerrado Papagaioin provincia MinasGeraes" (holotype, P, n.v.; Field Museumneg. 36982of probableholotype, photo at MICH). Eugenia ibitipocensis Cambessedes,in Saint-Hilaire,Flora Brasiliaemeridionalis2: 365. 1833. Type. Saint-Hilaires.n., "In praeruptusmontis alti vulgo Serra da Ibitipocain provincia MinasGeraes" (holotype, P, n.v.; Field Museumneg. 36969of probableholotype, photo at MICH). Eugenia expallens Berg, in Mart.Fl. Bras. 14(1):260. 1857.Type. Sellow s.n., "Habitatin prov. S. Pauli" (holotype, B, probablylost; Field Museumneg. 36952 of probableisotype at P, photo at MICH;Field Museumneg. 31568of probableisotype at W, photo at MICH). ?Eugeniaanceps Berg, in Mart.Fl. Bras. 14(1):260. 1857.Type. Sellow s.n., "Habitatin montibus Serra de Piedade prov. Minarum,""v. in hb. Kunth" (holotype, probablyB, apparently lost). Eugenia brachymischa Kiaerskou, Enum. Myrt. bras.: 165. 1893. Type. Glaziou 17003, "Rio"

(holotype, C; Field Museumneg. 20933,photo MICH).

Eugenia brachymischa forma pedunculata Kiaerskou, Enum. Myrt. bras.: 166. 1893. Type. Gla-

ziou 13891, Nova Friburgo(holotype, C; isotypes, F, RB). Luma bracteosa (A. P. de Candolle)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 528. 1941. Luma expallens (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 528. 1941. MyrceugeniacambessedeanaLegrandet Kausel,in Legrand,Comun.Bot. Mus. Hist. Nat. Montevideo 2(28):4. 1953.Illegitimatename because Eugenia brachymischaKiaerskouis cited as a synonym. The name Myrceugenia cambessedeana was proposed, based on Eugenia mon-

tana Cambessedes,because the name Myrceugeniamontana Kausel alreadyexisted.

Flora Neotropica

82 50

' 30

Sm;',,i''t_

;L;i'''

*? ,.{N-^^ .?I>^''<'*/'^ ~~

~~~~~~~~~~

Montevideo 2(28): 7. 1953.

*NM.

colchag;agt;'

FIG. 17.

''1'

Distributions of Myrceugenia colchaguensis and M. alpigena.

Myrceugenia bracteosa var. ibitipocensis (Cambessedes) Legrand, Comun. Bot. Mus. Hist. Nat.

Montevideo2(28):7. 1953. bracteosa var. expallens(Berg)Legrand,Comun.Bot. Mus. Hist. Nat. Montevideo Myvrceugenia 2(28): 6. 1953.

Myrceugeniamonticola Legrandet Kausel, in Legrand,Darwiniana11(2):321. 1957.Illegitimate name because Eugenia brachymischaKiaerskouis cited as a synonym. The name Myrceugenia monticola was proposed, based on Eugenia montana Cambessedes, because subsequent to the publicationof Myrceugeniacambessedeana(1953)the authorshad erroneously concludedthat the latter name was invalidatedby the earlierLuma cambessedeana (Berg) Burret(1941) [=Myrceugeniaglaucescens (Cambessedes)Legrandet Kausel].

Shrub or small tree 1-3 m high; hairs yellowish, golden-yellow, greyish-white to reddish-brown, a mixture of simple and dibrachiate, somewhat curled and matted together; twigs densely pubescent or tomentose when young, glabrescent with age, the bark light grey to reddish-brown; leaves elliptic, less often ovate, lanceolate, oblanceolate, or obovate, 2-6.5 cm long, 0.5-3.5 cm wide, 1.8-4.5 times as long as wide, densely pubescent to glabrous above, densely pubescent

below when young, losing some pubescence with age, the margin sometimes

Myrceugenia

slightly sinuate, sometimes revolute; apex acute or less often rounded, the midvein sometimes excurrent; base acute or cuneate; petiole channeled, apparently less so with age, 2-10 mm long, 0.5-2 mm thick, densely pubescent when young, glabrescent with age; midvein impressed above, prominentbelow; lateral veins indistinctor up to ca. 15 pairs, clearly visible; marginalveins indistinctto barely visible; blades light yellow-green or grey-greento dark olive-green above, often tinged with reddish-brown,lighterbut about the same colors below, coriaceous, sometimes strongly so, the upper surface sometimes lustrous; peduncles uniflorous, flattened, 0.5-5(-10) mm long, 0.5-1.5 mm wide, densely pubescent, solitary or up to 4 in a row in the axils of the leaves; bracteoles ovate, 1.9-4.5 mm long, 1-2.8 mm wide, 1.3-2 times as long as wide, densely pubescent to glabrous, or densely pubescent distally and glabrousproximallywithin, densely pubescent without, clasping the hypanthium;calyx-lobes ovate to oblong-ovate, 2-4 mm long, 1.5-3.8 mm wide, 0.8-1.7 times as long as wide, densely pubescent over entire inner surface or glabrous proximally, densely pubescent without; hypanthium densely pubescent, ca. 1-3 mm long; disk densely pubescent to glabrous, ca. 2-3 mm across; style ca. 5-7 mm long, sparsely pubescent to glabrous;stamens 80-220, ca. 7 mm long; anthers ca. 0.3-0.5 mm long when dry; petals glabrousor nearly so, ca. 2-4 mm in diam.; ovary generally 3-locular;ovules 71 per locule; fruit globose, ca. 8 mm in diam. Type. Martius s.n., "in Brasiliae Prov. Minarum," "(v. s. in h. Mart. sine nom." (holotype, M; Field Museum neg. 19920of holotype, photo at MICH). Specimens examined. BRAZIL. Minas Gerais: Serra da Piedade, Warming s.n. (C), Lund II 39 (C); south side of Serra da Piedade, ca. 5 km N of Caete, ca. 1800 m, 19 Jan 1971, Irwin et al. 28762 (MICH); Itamonte, V. Grande, 28 Apr 1962, Sobrinho 505 (F, MVM); prope Ibitipoca, Apr 1897, Magalhdes s.n. (R); Delfim Moreira, a 7 km de Is6polis, na estrada para Sao Francisco, 14 Oct 1968, Mattos 15386 (SP). Rio de Janeiro: Itatiaia, road to Agulhas Negras, km 8, ca. 2300 m, 18 Oct 1977, Landrum 2111 (MICH, R). Sio Paulo: Campos do Jordao, Parque Estadual, na Pedra Chorona, 2000 m, 25 Oct 1974, Mattos 15905 (SP); Campos do Jordao, Parque Estadual, 23 Apr 1974, Mattos 15800 (S); Campos do Jordao, hillside north of Capivari (ca. 22?45'S, 45?30'W), ca. 1600 m, 4 Dec 1977, Landrum 2798 (MICH, SP); Campos do Jordao, Reserva Florestal (ca. 22?45'S, 45?30'W), 1600 m, 5 Dec 1977, Landrum 2817, 2819, 2824, 2825, 2831 (at MICH, SP).

Myrceugenia bracteosa has been confused with M. alpigena var. rufa and the

differences between them are discussed under the latter species. Legrand and Kausel have recognized a separate species that includes the types of Eugenia montana, E. brachymischa and E. brachymischa forma pedunculata and have needlessly proposed two new names for it (M. cambessedeana and M. monti-

cola). If recognized as a separate species the epithet brachymischa should be taken up. These three type collections share leaves that are under 3 cm long and 1 cm wide and that have revolute margins, characteristicswhich suggest adaptation to a xeric habitat. As more collections are made, this group may emerge as a good taxonomic unit but I doubt that it will warrantspecific status. Myrceugenia bracteosa seems to extend no further south than Campos do Jordaoin Sao Paulo. A map of its distributionis shown in Fig. 18. Its apparently close relative, M. euosma, with which it is compared in the key, ranges from Paranasouth to Uruguay. Myrceugeniabracteosa flowersfrom Januaryto March and the fruits probablymaturein about October or November. 28. Myrceugenia euosma (Berg) Legrand, Anales Mus. Nac. Montevideo 4: 40. 1936. Figs. 7D, 7H, 18. Eugenia euosma Berg, Linnaea 27: 163 (name). 1856; in Mart. Fl. Bras. 14(1): 233. 1857. Eugenia euosma [var.] a lutescens Berg, in Mart. Fl. Bras. 14(1): 233. 1857. Type. Sellow s.n.

Flora Neotropica (holotype, B, apparentlylost; isotype MICH ex G designatedas lectotype). (Because the type of this taxon has been chosen as a lectotype of E. euosma, the namewould be replaced by E. euosma var. euosma.) Eugenia euosma [var.] f3 rufescens Berg, in Mart. Fl. Bras. 14(1): 233. 1857. Type. Sellow s.n.,

(holotype, B, apparentlylost; probableisotype, MICHex G). Eugenia nana Berg, Linnaea 27: 167 (name). 1856;in Mart. Fl. Bras. 14(1): 244. 1857. Type. Sellow s.n., "Habitatin Brasiliaeprov. St. Catharina"(holotype assumed to be the same as the holotype of E. nana [var.] a effusa, B, apparentlylost; probable isotype, MICH ex G). Eugenia nana [var.] a effusa Berg, in Mart. Fl. Bras. 14(1): 244. 1857. Type. Sellow s.n.,

(holotype, B, apparentlylost; probableisotype, MICHex G). (Because the type of this taxon is assumed to be the same as the type of E. nana, the name would be replacedby E. nana var. nana.)

?Eugenianana [var.] /3 congesta Berg, in Mart. Fl. Bras. 14(1): 244. 1857. Type. Sellow s.n. (holotype, B, apparentlylost). ?Eugeniaaprica Berg, in Mart.Fl. Bras. 14(1):218. 1857.Type. Sellow s.n., "Habitatad Campo do Cavalleiro,in Montevideo,floretJanuario"(holotype, B, apparentlylost). Luma euosma (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 534. 1941. Luma nana (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 535. 1941. Luma cinerea Burret, Notizbl. Bot. Gart. Berlin-Dahlem15: 534. 1941. Type. Fiebrig 6472 "Paraguay:In regione fluminisalto Parana"(holotype, presumablyB, apparentlylost; isotypes, LIL, SI). ?Lumaaprica (Berg) Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 535. 1941. Myrceugenia euosma var. oblongata Mattos, Loefgrenia 65: 5. 1975. Type. Hatschbach 20598

"Estado do Parana:Laranjeirasdo Sul" (holotype, MBM).

Tree or shrub up to 10 m high; hairs yellowish to reddish-brown,simple to symmetrically or asymmetrically dibrachiate; twigs densely pubescent when young, glabrescentwith age; leaves densely covered with appressedhairs below, puberulentabove, glabrescent with age above and below, elliptic, elliptic-oval, ovate, lanceolate to linear, 0.8-3(-4.7) cm long, 0.2-1.2(-1.5) cm wide, (1.7-)26 times as long as wide; apex acute to rounded;base cuneate to rounded;petiole strongly to slightly channeled, densely pubescent when young, glabrescentwith age, 1-3 mm long, 0.5-1 mm thick; midvein impressed or not above, prominent below; lateral veins indistinct, or up to about 15 pairs barely visible; marginal veins equallinglaterals in prominence;blades brown, yellow-greenor grey-green above, paler yellow-brown,tan or grey-greenbelow, membranousto coriaceous, the upper surface usually dull, less often lustrous; peduncles uniflorous(rarely bearing a triflorous dichasium), slightly to strongly flattened, 0.4-1.2 cm long, 0.5-1 mmthick, densely pubescent, solitaryor in pairs in the leaf axils; bracteoles ovate to lanceolate, 1.8-4.5 mm long, 1.2-2.5 mm wide, 1.2-2.3 times as long as wide, membranousto subcoriaceous,claspingthe hypanthium,densely pubescent without, pubescent or glabrouswithin; calyx-lobes triangularto ovate, 1.6-4 mm long, 1.3-3.3 mm wide, 0.9-1.6(-1.9) times as long as wide, subcoriaceous,densely pubescent without, densely pubescent over entire inner surface or only distally within; hypanthium densely pubescent, 1-2 mm long; disk 1.5-2 mm across, pubescent; style 5-8 mm long, sparsely pubescent; stamens 130-220, 4-10 mm long; anthers 0.3-0.5 mm long when dry; petals orbicularto slightly elongate, concave, 2.5-3.5 mm in diameter;ovary 2-4-locular; ovules (5-)7-15 per locule; fruit darkgrey, pubescent, 3-5 mm in diam.; seeds usually 2-3 per fruit, reniform to round, black 3-4 mm long. Type. Sellow s.n., type of Eugenia euosma [var.] or lutescens Berg hereby

designated as lectotype of Eugenia euosma Berg (lectotype MICH ex G).

Specimens examined. BRAZIL. Parana:Mun. CampoLargo, Bateiras, 18 Nov 1963, Hatschbach 10648(F, HB, LP, MBM, MVM);Mun. CampoTenente, Ribeiraoda Fazenda, 25 Jan 1968,Hatschbach 18446 (MBM, MVM, UC); Mun. Gal. Carneiro,13 Mar 1966, Hatschbach 15390(MBM, MO, MVM);Mun. Colombo,Bacaitava,30 Oct 1961,Hatschbach8780 (MBM, MVM);Mun.Clevelandia,

Myrceugenia

Varzea,900-1000m, 29 Dec 1956,Smith,Reitz & Caldato9570(HBR, MVM,R, US); Mun.Curitiba, Pinhaes, 22 Jan 1946, Hatschbach 171 (LIL, MBM, MICH, MVM);Mun. Curitiba,Bariqui, 1 Feb 1971, Hatschbach26205 (C, HB, MBM);Mun. Curitiba,900 m, 15 Sep 1914,Jonsson 931a (M, H); Mun. Contenda, near Contenda (ca. 25?40'S, 49?45'W), ca. 900 m, 7 Nov 1977, Landrum 2431, 2433,

2434 (MBM, MICH); Mun. Araucaria,rio Iguacu (ca. 25?35'S,49?30'W),ca. 900 m, 7 Nov 1977, Landrum2458 (MBM, MICH);Mun.Guarapuava,FazendaCampoReal, 1000m, 16 Dec 1965,Reitz & Klein 17790(HBR, MICH, MVM, NY, US); Mun. Guarapuava,Lagoa Seca, 6 Sep 1963,Pereira 7691 (HB, LP, MBM, MICH, MVM, RB); Mun. Guarapuava, Palmeirinha, 22 Oct 1960, Hatschbach

7355 (HB, HBR, MBM, MICH, MVM);Mun. Lapa, Nov 1957, Braga & Moreira507 (MVM, SP); Mun. Lapa, cerca de 15 km de Lapa, 5 Nov 1964, J. Mattos 11915 (SP); Mun. Lapa, 12-15 km southwest of Lapa (ca. 25?48'S, 49?44'W), 900-1000 m, 22 Jan 1965, Smith & Klein 14971 (HBR,

MICH, MVM, NY, R, US); Mun. Lapa, Serrinha, 17 Jun 1951, Hatschbach 2287 (MBM, MVM); Mun. Laranjeirasdo Sul, Laranjeirasdo Sul, 6 Nov 1963, Hatschbach 10337(F, HB, MBM, MVM, RB); Mun. Marmeleiro,estradaMarmeleiro-Campo Ere, 21 Feb 1971, Hatschbach26403 (C, MBM); Mun. Piraquara,Col. S. Roque, 6 Feb 1974,Hatschbach26304 (MBM, UC); Mun. Rio Negro, Doce Grande, 29 Nov 1956, Hatschbach3418 (HBR, MBM, MICH, MVM);Mun. Sao Joao do Triunfo, Sao Joao do Triunfo, 7 Nov 1967, Hatschbach 17758 (C, F, MBM, MICH, MVM, RB, UC, US), Hatschbach 17718 (C, F, MBM, MICH, MVM, RB, UC); Mun. Sao Mateus do Sul, Foz do Rio Taquaral,26 Oct 1956,Hatschbach3410 (MBM, MICH, MVM);Mun. Sao Mateusdo Sul, arredores da cidade, 14 Dec 1956, Hatschbach3576 (MBM, MVM);Mun. Timoneira,Timoneira,16 Jan 1957, Hatschbach 3581 (MBM, MICH, MVM); Mun. Tijucas do Sul, Vossoroca, Rio Sao Joaozinho, 15 Feb 1974, Kummrow355 (LP, MBM, MO); without specific locality, 21 Jan 1904, Dusen 3318 (R, US). Santa Catarina:Mun. AbelardoLuz, 12 km northof AbelardoLuz (ca. 26?32'S,52?20'W),9001000m, 23 Oct 1964,Smith & Reitz 12844(HBR, LP, MO, MVM,R, US); Mun. Bor Jardim,Morro Trombudo, 1500 m, 12 Dec 1958, Reitz & Klein 7834 (H, HBR, MVM, NY, UC, US); Mun. Bom Jardim,Serra do Orat6rio, 1400m, 12 Jan 1959, Reitz & Klein 8131 (H, HBR, MVM);Mun. Bom Jardim,1400m, 26 Jan 1971,Eskuche01471(CTES, MVM),ca. 1250m, 27 Jan 1971,Eskuche1647b4 (CTES); Mun. Bom Retiro, Riosinho, 1000m, 24 Dec 1948, Reitz 2770 (H, HBR, MVM), 21 Dec 1948,Reitz3651 (LIL, PACA, W); Mun. Bor Retiro, Campodos Padres,2000m, 16 Dec 1948,Reitz 2420 (HBR, MVM, PACA, R), 1900m, 17 Dec 1948, Reitz 2426 (H, HBR, MVM), Reitz 3697 (H, US), 1650 m, 17-19 Nov 1956, Smith, Reitz & Klein 7688 (HBR, MVM, NY, R, US); Mun. Bor Retiro, Reitz 1978 (HBR, MICH, MVM), 900 m, 9 Jan 1948, Reitz 3577 (F, NY, UC, W); Mun. Campo Alegre, fazenda of Ernesto Scheide, 900-1100 m, 9 Nov 1956, Smith & Klein 7524 (HBR, MVM, R, UC, US), 1 Feb 1957, Smith & Klein 10557 (HBR, MVM, R, RB, US), Smith & Klein

10564(HBR, MVM, NY, R, US); Mun. Canoinhas,BarraGrande,750 m, 11 Dec 1962, Klein 3761 (HBR, MVM, NY); Mun. Canoinhas,west of Canoinhason road to P6rto Uniao, ca. 750 m, 17 Dec 1956, Smith & Reitz 8573 (RB); Mun. Cacador,8 km north of Ca9ador,950-1100 m, 21 Dec 1956, Smith & Reitz 8970 (MVM, US); Mun. Chapec6, 24 km west of Campo Ere, 900-1000 m, 20 Feb 1957, Smith, Reitz & Sufridini9484 (HBR, MVM, R, US); Mun. Chapec6,9 km west of CampoEre, 900-1000 m, 20 Feb 1957, Smith & Klein 11540 (HBR, MVM, R, US); Mun. Curitibanos,6 km northwest of Lebon Regis, 700-900 m, 8 Feb 1957, Smith & Klein 11043 (HBR, MVM, R, US); Mun.

Curitibanos,northof Curitibanos,850-950 m, 6 Dec 1956,Smith & Klein8369 (HBR, MICH,MVM, R, US); Mun. Dionisio Cerqueira,near Dionisio Cerqueira,800-850 m, 30 Dec 1956, Smith & Klein 9649 (HBR, MVM, R, US); Mun. Guaraciaba,13 km northwestof Sao Migueld'Oeste (ca. 26?32'S, 53?34'W), 400-600 m, 19 Dec 1956, Smith & Klein 14156 (GH, HBR, MICH, MVM, NY, UC, US);

Mun. Joacaba, 15 km east of Ponte Serrada,700-900 m, 3 Jan 1957, Smith & Reitz 9877 (HBR, MVM, R, US); Mun. Bom Jardim,Fazenda da Laranja,1400m, 10 Dec 1958, Reitz & Klein 7741 (H, HBR, MVM, NY, UC, US); Mun. Lajes, na base do MorroTributo, 26 Dec 1956, J. Mattos 3916 (C); Mun. Lajes, Indios, 950 m, 16 Dec 1967, Lourteig2266 (C, CTES, MVM);Mun. Lajes, 23 km north of Lajes, 800-900 m, 4 Dec 1956, Smith & Klein 8231 (HBR, MVM, NY, R, US); Mun. Lajes, near Passo de Socorro, elev. 800-900 m, 14 Jan 1957, Smith & Reitz 9943 (HBR, MVM, R, US); Mun. Lajes, between Palmeirasand Lajes, 800-900 m, 2 Dec 1956, Smith & Klein 8113 (HBR, MICH, MVM, R, US); Mun. Lajes, ca. 15 km east of OtacilioCosta (ca. 27?25'S,50?W),ca. 1000m, 21 Nov 1977,Landrum2648, 2653 (MBM,MICH);Mun.Lajes, ca. 18km south of Lajes (ca. 27?50'S, 50?20'W),ca. 1000m, 22 Nov 1977,Landrum2680 (MBM, MICH);Mun. Mafra,5 Dec 1948,Hatschbach 1132 (MBM, MICH, MVM); Mun. Mafra, Campo Novo, 800 m, 7 Sep 1957, Reitz & Klein 4912

(HBR, MICH, MVM, US); Mun. Mafra,11 km east of Mafraon the roadto Tingui,ca. 800 m, 8 Dec 1956, Smith & Klein8468 (HBR, MVM, R, US); Mun. Mafra,7-9 km west of Tingui, 800 m, 2 Feb 1957, Smith & Klein 10614 (HBR, MVM, R, US); Mun. Rio Negrinho, 5-14 km west of Rio Negrinho, ca. 800 m, 2 Feb 1957, Smith & Klein 10606 (HBR, MICH, MVM, R, RB, US); Mun. Rio Negrinho, 24 Jan 1953, Reitz 5192 (H, HBR, MVM, PACA, US, W); Mun. Mafra, ca. 15 km west of Rio Negrinho (ca. 26?15'S, 49?40'W), ca. 900 m, 24 Nov 1977, Landrum 2712, 2717, 2718, 2719 (MBM,

MICH);Mun. Bom Jardim,1 km east of Bom Jardimda Serra,elev. 1100-1200m, 16Jan 1957,Smith

Flora Neotropica

& Reitz 10195(HBR, MVM, R, US); Mun. Sio Jaoquim,3-7 km west of the Serrado Orat6rio,1200 m, 16 Jan 1957, Smith & Reitz 10178 (HBR, MVM, R, US); Mun. Iriane6polis,Valoes, 750 m, 10 Dec 1962, Klein 3740 (HBR, MBM, MVM, US); Mun. Xanxere, 8 km northof AbelardoLuz, 500600 m, 19 Feb 1957, Smith & Klein 11469 (HBR, MVM, R, US); Mun. Lajes, 10 Jan 1951, Rambo

49649 (LIL, PACA);Tres Barras,25 Feb 1948,A. Mattos & Labouriaus.n. (MVM,RB). Rio Grande do Sul: Caxias, Piai, 21 Jun 1950, Rambo 47189 (H, MVM, PACA); Sao Francisco de Paula, Vila Oliva, 30 Dec 1946, Rambo30795 (F, MVM, PACA, W); Taimbesinhno,18 Dec 1950,Rambo49372 (HBR, LIL); Mun. Sao Francisco de Paula, cascatinhanear Sao Francisco de Paula (ca. 29?30'S, 50?15'W),ca. 1000m, 26 Dec 1977, Landrum2965 (MICH, PACA); Mun. Cambara,near Cambara (ca. 29?S, 50?W),ca. 1000 m, 27 Dec 1977, Landrum2986 (MICH, PACA); Mun. Rolante, Santa Tereza, Fazenda Englert (ca. 29?25'S,50?20'W),ca. 1000 m, 28 Dec 1977, Landrum3005 (MICH, PACA); Montenegro,S. Salvador, 10 Dec 1946, Henz s.n. (LIL); Mun. Soledade, Farqueta,23 Jan 1964, Pereira 8565 (HB, LP, MVM, RB); Soledade, 13 Feb 1951, Rambo 50046 (LIL); Passo do Socorro p. Vacaria,27 Dec 1951, Rambo51648 (MVM, PACA, US). PARAGUAY.Regi6ndel Guaira,Paso Yobay, Apr 1938,Rojas 7901(H, MVM);regi6ndel Guaira, San Augustin,Jun 1932, Rojas 6054 (MVM);region Caaguazf, Villa Pastoreo, Jun 1932, Rojas 5995 Feb 1972,Schinini4358 (CTES, SI); Piribebuy-Piraretah, (MVM);Paraguari,SaltoPirareta-Piribebuy, Mar 1942, Rojas 9609 (H).

URUGUAY. CerroLargo,25 Feb 1938,RosengurttB2582(H, MVM);CerroLargo,FrayleMuerto, Azotea PadreAlonso, 19 Mar 1945,RosengurttB4801/2(MICH),RosengurttB4801(A, LIL, MICH, MVM, SI, SP, US); CerroLargo, CerroGuazunambu,Feb 1928, Schroeders.n. (H, MVM);Rivera, Alrededoresde la Ciudad,22 Jan 1944, Legrand3480 (MICH, NY); Rivera, Ataques, 14 Jan 1941, Legrand2492 (H, MICH, NY); Tacuaremb6,Grutade los Helechos, Jan 1940, Legrand2117 (H, MICH, MVM, NY); Treintay Tres, Serraniasdel Yerbal, Apr 1936, Legrand684, 734 (H, MVM); Treintay Tres, Quebradade los Cuervos, 1 Apr 1936, Legrand754 (F, H, MICH, MVM, NY). ARGENTINA. Misiones: Dep. San Javier, Panambi, 23 Jan 1947, Schwarz 3958 (MVM); San Antonio NE de Misiones, 2 Nov 1944,Bertoni470 (NY).

Eugenia nana [var.] f congesta Berg is probably conspecific with Eugenia nana [var.] a effusa and therefore is included as a possible synonym. Burret referred Eugenia aprica to Luma and he probably saw the type. Most of what Burretthoughtto be Luma is now consideredto be Myrceugenia.The description of E. aprica fits M. euosma well and it was collected within the range of M. euosma. Thereforeit also considered to be a possible synonym. Myrceugenia euosma is one of the most xerophytic species of Myrceugenia,

and is found from the Araucaria forests of the planalto south to Uruguayand as far west as Paraguay.A map of its distributionis shown in Fig. 18. In Paraguay it sometimes appears quite similarto M. glaucescens. Myrceugenia euosma is similar to M. bracteosa and is compared to that species

in Key I. It is also compared with M. alpigena var. alpigena in the discussion of that species. There are two specimens collected by Glaziou (19366, 19369)at C which have "St. Paul" and "S. Paul" on the labels. Althoughthis probablymeans Sao Paulo, it is unlikely that these specimens actually come from the state of Sao Paulo because there are no other known collections from that state. Otherwise, as far as is known, Myrceugeniaeuosma grows no farthernorththan Parana.It flowers from November to Februaryand the fruits seem to matureabout 1 year later. Certain plants in Santa Catarinaand Parana appear to be hybrids between Myrceugenia euosma and M. glaucescens. They are discussed in the section on hybridization.

29. Myrceugeniaacutiflora(Kiaerskou)Legrandet Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 5. 1953. Figs. 19, 33A. Eugenia acutiflora Kiaerskou,Enum. Myrt. bras.: 164. 1893. Luma acutiflora(Kiaerskou)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 528. 1941.

Myrceugenia 60

$. 0*e. FIG. 18.

frianctocensis a"'

Distributions of Myrceugenia franciscensis, M. bracteosa, and M. euosma.

Shrub or tree 2-12 m high; hairs reddish-brown, dibrachiate, mainly symmetrical; twigs densely pubescent when young, glabrescent with age, the bark reddish-brown to grey, often becoming a mass of thin papery scales; leaves elliptic to ovate, 2.5-4.5 cm long, 1-2 cm wide, 1.9-2.9 times as long as wide, the upper surface entirely glabrous or sparsely pubescent along the midvein, the lower surface densely pubescent when young, losing some or nearly all pubescence with age; apex acute to acuminate, usually bluntly so, rarely rounded; base acuminate, cuneate or acute; petiole strongly channeled, densely pubescent, 23 mm long, 0.5-1 mm thick; midvein impressed proximally or for entire length above, prominent below; lateral veins indistinct or up to ca. 15 pairs faintly

Flora Neotropica

visible; marginal veins equalling lateral or less distinct; blades yellow-green, grey-green, or reddish-brown above, about the same color but lighter below, coriaceous, the upper surface sometimes lustrous; peduncles uniflorous, flattened, 3-10 mm long, 1-1.5 mm wide, densely pubescent, solitary or two in a row in the leaf axils; bracteoles ovate to lanceolate, 2.5-4.5 mm long, 1.5-2 mm wide, 1.5-2.5 times as long as wide, subcoriaceous, sparsely pubescent to glabrous within, densely pubescent without, clasping the hypanthium; calyx-lobes ovate to triangular, 3-5.5 mm long, 2.5-3.5 mm wide, 1-1.7 times as long as wide, densely pubescent over entire inner surface or less densely pubescent proximally, densely pubescent without; hypanthium obconic, densely pubescent, 2-3 mm long; disk sparsely pubescent, ca. 3 mm across; style 6-8 mm long, very sparsely pubescent to glabrous; stamens 160-230, 3-8 mm long; anthers 0.3-0.6 mm long when dry; petals densely pubescent without, glabrous to very sparsely pubescent within, 3-4 mm in diam.; ovary 3-4-locular; ovules 8-14 per locule; fruit globose, ca. 7-10 mm in diam.; seeds ca. 4-7. Types. Glaziou 13894, hereby designated as lectotype (lectotype, C; isolectotypes, F, R, RB, US; Field Museum neg. 23509 of lectotype, photo at MICH); Glaziou 13888 (paratype, C). Specimens examined. BRAZIL. Rio de Janeiro: Tijuca, 22 Feb 1927, Pessoal do Horto Florestal s.n. (MBM, NY, RB-752); Tijuca, on trail to Bico do Papagaio, 800 m, 24 Oct 1977, Landrum 2205 (MICH, RB), 1000 m, 24 Oct 1977, Landrum 2192 (MICH, RB); Sumare, 24 Feb 1964, Pereira & Duarte 4522 (HB, HBR, MICH, RB); Parque Nacional da Serra dos Orgaos, Terez6polis, 29 Jun 1942, Dionisio & Otario 216 (R). Parana: Bocaiuva, 29 Mar 1941, Cecato 16 (MICH, RB); Mun. Quatro Barras Rancho Velho, 23 Feb 1964, Hatschbach 10995 (F, HB, MBM, MICH, MVM); Mun. Sao Jose do Pinhais, road to Guaricana (ca. 25?40'S, 49?W), 900-1000 m, 4 Nov 1977, Landrum 2393, 2407 (at MBM, MICH), 15 Dec 1977, Landrum 2906, 2931 (at MBM, MICH). Santa Catarina: Mun. Bom Retiro, Paulo Lopes, 400 m, 24 Aug 1973, Bresolin 801 (MVM); Mun. Florian6polis, Morro Costa da Lagoa, 16 Mar 1967, Klein & Souza 7324 (MVM), 300 m, 13 Feb 1969, Klein & Bresolin 8173 (HBR, MVM), 13 Mar 1969, Klein 8255 (HBR, MVM), Mun. Gov. Celso Ramos, Jordao, 250 m, 11 Aug 1971, Klein 9648 (MVM).

Myrceugenia acutiflora is most similar to M. myrcioides var. myrcioides and M. pilotantha var. major. From the first it can be distinguished by its calyx-lobes that are not valvate in the bud nor concave throughout the inner surface. The leaf apex is usually a good indicator as well, normally being bluntly acuminate in M. acutiflora and sharply acuminate in M. myrcioides. Myrceugenia acutiflora can be distinguished from M. pilotantha var. major by its hairs that are all dibrachiate and by its petiole which is strongly channeled. In M. pilotantha var. major the hairs are a mixture of simple and dibrachiate and the petioles are unchanneled or only scarcely channeled. Myrceugenia acutiflora flowers, as far as is known, from January to March and probably fruits in the winter months of July and August. Myrceugenia myrcioides var. myrcioides has its flowering peak from April to May and therefore phenology also serves to separate these entities. Myrceugenia acutiflora seems to be confined to higher elevations along the Atlantic coast from Rio de Janeiro to Santa Catarina. It is an understory shrub or tree of the misty wet forests of this region. A map of its distribution is shown in Fig. 19. 30. Myrceugenia venosa Legrand, Darwiniana 11(2): 323. 1957.

Fig. 19.

Myrceugenia miersiana var. lanceolata Legrand, in Legrand et Klein, Flora Ilustr. Catarinense [Mirt]: 401. 1970. Type. Klein 1925, Santa Catarina, Ibirama, Horto Florestal do I. N. P., 350 m, 7 Mar 1956 (holotype, MVM; isotypes, H, HBR, US).

Myrceugenia 50

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Small tree or shrub, 2-7 m high; hairs reddish-brown to yellowish-brown, symmetrically dibrachiate to simple, mainly dibrachiate on the leaves and flowers, a mixture of simple and dibrachiate on the twigs, appressed to slightly spreading; twigs densely pubescent when young, glabrescent with age, the bark of older twigs light grey to tan, remaining more or less smooth; leaves elliptic to oblanceolate, 5-10 cm long, 1-3 cm wide, 2.3-4.6 times as long as wide, the upper surface glabrous or with scattered hairs on the blade, sparsely to densely pubescent along the midvein, the lower surface moderately pubescent when young, usually slightly excurrent; base acute, cuneate to barely acuminate; petiole channeled, densely pubescent, 2-6 mm long, 1-1.5 mm thick; midvein impressed for entire length above, quite prominent below; lateral veins sometimes slightly impressed above, 10-15 pairs prominent below, other less prominent secondary veins visible between these; marginal veins prominent, arching slightly between the stronger laterals, a much weaker secondary lateral, exterior to the main lateral

Flora Neotropica

sometimes visible; blades coriaceous, dull to slightly lustrous, darkor light greygreen to yellow-green(often tingedwith reddish-brownupon drying)above, about the same color but lighterbelow; peduncles uniflorous, 1-1.5 mm long, ca. 1 mm wide, densely pubescent, solitary or up to 3 in a row in the leaf axils; bracteoles ovate to lanceolate, 2.5-4 mm long, 1.5-3 mm wide, 1.3-2.2 times as long as wide, subcoriaceous, densely pubescent without, sparsely to densely pubescent within, clasping the hypanthium; calyx-lobes ovate to triangular,overlapping slightly, sometimes appearingto be valvate in the bud, coriaceous, 2.5-5.5 mm long, 2-4.5 mm wide, 1-1.5 times as long as wide, densely pubescent without, densely to sparsely pubescent within; hypanthiumobconic, densely pubescent, 2-3 mm long; disk 2.5-4 mm across, sparsely pubescent; style ca. 5-9 mm long, sparsely pubescent; stamens ca. 250-400, ca. 5-9 mm long; anthers 0.3-0.5 mm long when dry; petals suborbicular,4-5 mm in diam., sparsely pubescent to glabrous within, densely pubescent without, the marginslightly irregular;ovary usually 3-locular;ovules 10-20 per locule; fruit ca. 1 cm in diam. Type. Reitz 5474, Santa Catarina,Serra da Boa Vista, ca. 1100 m (holotype, MVM; isotypes, H, HBR, US). Specimens examined. BRAZIL. Parana:Mun. Piraquara,Novo Tirol, 26 Apr 1964, Hatschbach 11342(HB, MBM, MICH, MVM);Mun. Sao Jose dos Pinhais,roadto Guaricana(ca. 25?40'S,49?W), 900-1000 m, 4 Nov 1977, Landrum 2409, 2410, 2411 (MBM, MICH), 15 Dec 1977, Landrum 2921,

2932, 2933 (MBM, MICH). Santa Catarina:Mun. Vidal Ramos, Sabia, 600 m, 31 Dec 1957, Reitz & Klein 5991 (H, HBR, MVM).

This species has been poorly understood.Legrandfirst described Myrceugenia

venosa and later described a second taxon, M. miersiana var. lanceolata, which

I believe to be a later synonym. In the protolog of the second taxon he states that it is apparentlya hybridbetween M. miersiana and M. mycioides, an opinion with which I concurred until I was able to observe all three species growing sympatricallyin Parana.There M. venosa was obviously morphologicallydistinct from both the other species and was floweringat least a month before them. Myrceugenia venosa is in some ways intermediate between M. myrcioides and

M. miersiana as can be seen in the following table. M. mvrcioides

Calyx-lobesvalvate Twig hairs dibrachiate Leaf hairs dibrachiate Lateral and marginal veins not normallyprominent

M. venosa

Calyx-lobes slightly imbricate, appearingvalvate Twig hairs mixed Leaf hairs dibrachiate Lateral and marginal veins prominent

M. miersiana

Calyx-lobesimbricate Twig hairs simple Leaf hairs simple Lateral and marginal veins prominent

Myrceugeniavenosa is distinctfrom the others in having leaves that are on the average longer and narrowerthan either, marginaland lateralveins that are even stronger than in M. miersiana and in having the lower surfaces of the leaves sparsely pubescent at maturity rather than densely pubescent as usual in M. myrcioides and M. miersiana. It also differs from M. miersiana in having a

channeled petiole and it can be distinguishedfrom M. myrcioides by the closed flower bud which is slightly pointed ratherthan an indented hemisphere. Myrceugenia venosa flowers from December to April and the fruits probably maturefrom about July to October. It seems to grow along the eastern edge of the planalto. A map of its distributionis shown in Fig. 19.

Myrceugenia

31. Myrceugeniamyrcioides(Cambessedes)Berg, Linnaea 27: 134. 1856. Eugenia myrcioidesCambessedesin Saint-Hilaire,Flora brasiliaemeridionalis2: 353. 1832.

Small tree 1.5-6 m high; hairs reddish-brownor yellowish-brownto greyishwhite, dibrachiate, symmetric or nearly so, usually appressed; twigs densely pubescent when young, glabrescentwith age, the bark sometimes falling in thin sheets; leaves usually elliptic, elliptic-ovate or lanceolate, less often elliptic-obovate or oblanceolate, 3.5-16 cm long, 1-7 cm wide, 1.8-3.6 times as long as wide, glabrousto sparsely pubescent above (or densely pubescent along the midvein), sparsely to densely pubescent below, losing some pubescence with age; apex usually acute or acuminate (sometimes abruptly so), less often rounded; base cuneate, acute, or acuminate;petiole channeled, densely pubescent, 3-12 mm long, 1-2 mm thick; midvein impressed for entire length or only proximally above, prominentbelow; lateral veins indistinctor up to ca. 30 pairs moderately prominent below; marginal veins equalling laterals or less prominent; blades coriaceous to membranous,dull light grey-green or yellow-green to dark greygreen above (often tinged with reddish-brownwhen dry), about the same color or a lighter shade below; peduncles uniflorous, 0.5-2.5(-4.5) cm long, 1-2 mm wide, rarely lacking, densely pubescent, often slightly sulcate, solitary or up to 4 in a row in the leaf axils; bracteoles ovate to lanceolate, 1.5-8 mm long, 0.94 mm wide, 1.3-5.5 times as long as wide, coriaceous to subcoriaceous, densely pubescent without, densely pubescent to sparsely pubescent within, usually clasping the hypanthium,persistent until the fruit matures or sometimes deciduous before the fruit maturesin var. acrophylla; calyx-lobes ovate to triangular, concave throughoutentire inner surface, valvate or overlappingalong the narrow, membranousmargins of two lobes, otherwise coriaceous, tightly closed in the bud, 2.5-6.5 mm long, 2-6 mm wide, 0.8-1.7 times as long as wide, densely pubescent to glabrous without and within; hypanthiumobconic, densely pubescent, 2-5 mm long; disk 2.5-6 mm across, sparsely to densely pubescent; style 5-10 mm long, sparsely pubescent to glabrous; stamens 130 to over 500, 5-17 mm long; anthers 0.4-0.6 mm long when dry; petals more or less orbicular,less often clawed, 2.5-7.5 mm in diam., sparselypubescent to glabrouswithin, densely pubescent to glabrous without, the margin sometimes irregular;ovary 3-4locular; ovules (5-)7-14 per locule;fruit purpleblack to grey, 1-1.5 cm in diam., puberulentto pubescent; seeds about 3-5, oblong, 0.6-1 mm long. The two varieties of Myrceugeniamycioides are undoubtedlyclosely related and togetherform a logical taxonomicunit. It is usually possible to separatethem using calyx-lobe length and the presence or absence of dense pubescence within the calyx-lobes. Leaf size, hair size and hair color are usually correlated with these characteristicsof the calyx-lobes. But because division is often difficult, especially in the northernpart of their range, and because the two entities form a naturalunit, they have been retainedas one species. When one compares the floweringperiods of the two varieties, it is apparent that they reach maxima at slightly dfifferenttimes of year, Myrceugenia myr-

cioides var. acrophylla from January to March, and M. myrcioides var. myr-

cioides in April and May. In the table below are shown all the phenologicaldata for the floweringspecimens availableto me, except for my own collections made in October throughDecember, which would have skewed the results towards the monthsin which I collected. There were a total of 43 collections of M. myrcioides var. acrophylla and 33 of M. myrcioides var. myrcioides.

Flora Neotropica

Months 2 1

3 9%

2 6%

1 3%

myrcioides

numberof collections % of total

1 3%

3 9%

5 2 8 8 6% 15% 24% 24%

acrophylla

numberof collections % of total

1 1 2 7 9 12 9 2% 2% 5% 16% 21% 28% 21%

2 5%

The differences in flowering periods are probably more pronounced than is indicatedin the table because collectors generally do not collect in proportionto the numberof floweringindividualsencountered. Whetherthe differencein flowering period is an adaptationfor maintainingreproductiveisolation or caused by some other factor is an interesting problem. The two varieties have nearly the same range and often grow near one another. Assumingthat they are adaptedto somewhat different niches, and that they are still capable of interbreeding,then it would be to their mutualadvantageto have differentfloweringperiods. The fruits of both varieties seem to maturemainly in the late winter and spring months from July to November. It is not obvious to me what ecological differences there are between them. They both grow in the planaltoand in the coastal lowlands. Distributionmaps of both varieties are shown in Fig. 19. Key to Varieties of Myrceugenia myrcioides 1. Calyx-lobes densely pubescent within and without, usually 2.5-4.2 mm long, 2-3.5 mm wide; stamensca. 130-220;leaves usually3.5-9 cm long, 1-4 cm wide;hairsusuallyreddishbrown to yellowish-brown, most usually surpassing 0.5 mm long; bracteoles 1.5-2.5 a. var. myrcioides. (-4) mm long. 1. Calyx-lobesentirelyglabrousto very sparselypubescentdistallywithin, densely pubescent to glabrouswithout, usually 4-6.5 mm long, 3.5-6 mm wide; stamens ca. 250 to over 500; leaves usually 4-16 cm long, 2-7 cm wide; hairs usually greyish-white,less often reddishbrown to yellowish-brown,often all less than 0.3 mm long; bracteoles(2-)3-8 mm long. b. var. acrophylla.

31a. Myrceugeniamyrcioidesvar. myrcioides

Figs. 19, 33B.

Eugenia myrcioidesCambessedesin Saint-Hilaire,Florabrasiliaemeridionalis2: 353, as to type, 1832. EugeniahypericifoliaGardner,LondonJ. Bot. 2: 354. 1843.Type. Gardner421, "Hab. In woods by sides of streams. Fl. March."Serrados Orgaos(holotype, K; Field Museumneg. 23547 of isotype at G, photo at MICH). Eugenia obtusiflora Kiaerskou, Enum. Myrt. bras.: 168. 1893. Type. Glaziou 13885, hereby designatedas lectotype (lectotype, C; isolectotype, US); Glaziou5867 (paratype,C). Luma myrcioides(Cambessedes)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 531. 1941. Luma obtusiflora(Kiaenskou)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 531. 1941. Myrceugeniamyrcioidesvar. hypericifolia(Gardner)Legrand,in Legrandet Klein, FloraIlustr. Catarinense[Mirt]:407. 1970. ?Mvrceugenia mvrcioides var. loefgreniana Mattos, Loefgrenia 71: 1. 1977. Type. A. Loefgren

s.n., "ex Comm. Geogr. Gebl. de Sao Paulo no. 3554," "Estado de Sao Paulo: C6rrego Alegre" (holotype, SP, n.v.).

Hairs reddish-brownto yellowish-brown, most usually over 0.5 mm long;

leaves 3.5-9 cm long, 1-4 cm wide; blades coriaceous; bracteoles 1.5-2.5(-4)

mm long, 0.9-1.7 mm wide, 1.3-2.7 times as long as wide, densely pubescent within and without, persistent until the fruit matures; calyx-lobes ovate to tri-

Myrceugenia

angular,2.5-4.5 mm long, 2-3.5 mm wide, valvate in the bud, coriaceous, densely pubescent within and without; stamens ca. 130-220. Type. Saint-Hilaire s.n., "In sylvis prope Sebastianopolim.Fructus maturat Septembri" (holotype, P, n.v.; Field Museum neg. 36986 of apparentholotype, photo at MICH). Specimensexamined.BRAZIL. MinasGerais:DelfimMoreira,a 7 km de Is6polis, na estradapara Sao Francisco, 14 Oct 1968, J. Mattos 15389(SP). Rio de Janeiro:estrada Vista Chinesa, caminho do encanamento,29 May 1963, Martins290 (CTES, GUA, MVM, SI); TijucaNationalParknearRio de Janeiro,near caves on road to Bom Retiro, 600-700 m, 12 Oct 1977, Landrum2004 (MICH,RB); matadas ObrasPtiblicas,perto da sede do Horto Florestal, 30 Apr 1927,Pessoal do Horto Florestal s.n. (MBM, MICH, RB, US); above Silvestre near Rio de Janeiro, 18 Jul 1927, Harshberger885 (US); Serrada Carioca, 13 Jun 1945, Occhioni256 (HBR, MICH, MVM, RB); Serrada Estrella, ca. 2 km above Meio da Serra near Petr6polis(ca. 22?30'S,43?W),ca. 500 m, 16 Oct 1977, Landrum 2067, 2073 (MICH, RB); CampoGrande,Serrado Mendanha,27 Apr 1958, Costa 9 (R). Sao Paulo: Parelheiro,perto de Santo Amaro, Sep 1961,Silveiras.n. (SP); Sales6polis, na Estacao Biologicade Boraceia,perto das margensdo Rio Coruja,27 Apr 1966,J. Mattos 13491(SP); Mongagua,ao pe da Serra do Mar, Mar 1964, J. Mattos 11807 (MBM, SP); ca. de 8 km ao nordeste de Juquia,4 Apr 1961, J. Mattos 8869 (C, SP). Parana:CampinaGrande do Sul, Sitio do Belizario, 7 Apr 1967, Hatschbach 16242(C, MBM, MO, MVM, NY, UC); Mun. Castro,Abapa,28 Feb 1961, Hatschbach 7763 (MBM, MVM); Mun. Curitiba,Santa Felicidade, 17 Feb 1929, Gurgel s.n. (RB); Mun. Gal. Carneiro,Cab. Rio Iratim, 18 Oct 1966, Hatschbach 15000 (MBM, MICH, MVM, UC, US); Mun. GuaraqueSaba,Serrinha,9 Aug 1967, Hatschbach 16896(C, MBM, MICH, MO, MVM, NY, UC), 10 Apr 1968, Hatschbach 19005 (HB, MBM, MICH, MO, MVM, NY, UC); Mun. Guaraquegaba, Rio Taga9aba,20-50 m, 20 May 1968, Hatschbach 19221(C, CTES, HB, MBM, MICH, MO, MVM, NY, SP, UC); Mun. Guaraquegaba,Rio do Cedro, 30-50 m, 8 Mar 1968, Hatschbach 18678 (C, CTES, MBM, MVM, NY, UC); Mun. Guaraquecaba,ca. 30 km towards Antoninafrom Guaraquecaba, ca. 200 m, 14 Dec 1977,Landrum2890 (MBM, MICH);Mun. Morretes,Pilao de Pedra, 22 Jul 1966, Hatschbach 8065 (HB, HBR, MBM, MICH, MVM); Mun. Piraquara,Roga Nova, 27 May 1951, Hatschbach2269 (MBM, MVM);Sio Jose dos Pinhais, Vossoroca, 15 Aug 1951, Hatschbach 2462 (MBM, MICH,MVM);Mun. Sio Jose dos Pinhais,Rio Pequeno,950 m, 3 Jun 1970,Hatschbach 24377 (C, CTES, HB, LP, MBM, MO, NY, UC); Mun. Sao Jose dos Pinhais,road to Guaricana(ca. 25?40'S, 49?W), 900-1000 m, 1 Nov 1977, Landrum 2305, 2312, 2318 (MBM, MICH), 15 Dec 1977, Landrum 2914, 2923, 2934 (MICH); Mun. Sio Jose dos Pinhais, Malhada, 26 Dec 1960, Hatschbach

7604 (MBM). SantaCatarina:Mun. Cacador,Rio dos Bugres, 800 m, 8 Jan 1962,Reitz & Klein 11750 (H, HBR, MVM);Mun. Cagador,MadeireiraRio Verde, 1000-1200m, 48 km west of Ca9ador,ca. 26?45'S, 51?22'W, 2 Dec 1964, Smith & Klein 13366 (HBR, MVM, R); Mun. Campo Alegre, Morro

Iquererim,1000-1300m, 9-10 Dec 1956, Smith & Klein 8499 (HBR, MVM, R, US); Mun. Campo Alegre, upper fazenda of Ernesto Scheide, 900-1100 m, 9 Nov 1956, Smith & Klein 7507 (HBR, MVM, R, RB, UC, US); Mun. Ibirama,Horto Florestal I. N. P., 350 m, 18 May 1956, Klein 1988 (H, HB, HBR, MBM, NY, US); Mun. Itajai, Morroda Ressacada, 150 m, 6 May 1955, Klein 1366 (H, HBR, MVM);Mun. Itajaf,BragoJoaquim,Luis Alves, 300 m, 22 Mar 1956, Reitz & Klein2875 (H, HB, HBR, MBM, MVM, NY, PACA, US); Mun. Itajaf, Morro da Fazenda, 100 m, 14 May 1954, Reitz & Klein 1837 (H, HBR, MVM, NY, UC), 150 m, 1 Jul 1954, Reitz & Klein 1901 (H, HB, HBR, MBM, MVM, NY, PACA, UC, US); Mun. Jacinto Machado, Sanga da Areia, 200 m, 4 Sep 1959, Reitz & Klein 9007 (H, HBR, MVM), 250 m, 13 May 1960, Reitz & Klein 9672 (H, HBR, MVM); Mun. Lajes, Alto da Serra, Encruzilhada, 900 m, 13 Sep 1962, Klein 2905 (H, HBR, MVM); Mun. Palhoga, Anitapolis, ca. 500 m, 4 Apr 1953, Klein 490 (H, HBR, MVM); Mun. Sao Jose, Serra da Boa Vista, 900 m, 11 Aug 1960, Reitz & Klein 9724 (H, HBR, MVM, NY), 800 m, 3 Mar 1961, Reitz & Klein 10835 (H, HBR, MVM), 700 m, 25 Jan 1961, Reitz & Klein 10756 (H, HB, HBR, MBM, MVM, US), 3 Mar 1961, Reitz & Klein 10846 (H, HBR, MVM, US); Mun. Sombrio, Garapuvu, Vista Alegre, 30 m, 14 May 1960, Reitz & Klein 9677 (H, HBR, MVM, NY, UC, US); Mun. Vidal Ramos, Sabia, 750 m, 16 Apr 1957, Reitz & Klein 4295 (GH, H, HBR, MVM, NY, US).

Myrceugenia myrcioides var. myrcioides is quite similar to M. acutiflora and

is probably derived from that species or a not too distant ancestor. Their dis-

tinction is discussed under M. acutiflora. Myrceugenia myrcioides var. myrcioides might also be confused with M. miersiana, M. pilotantha and M. venosa.

The first two species can be distinguishedfrom M. myrcioides var. myrcioides by their imbricatecalyx-lobes and simple hairs. Myrceugeniavenosa has strong lateral and marginalveins which M. myrcioides var. myrcioides does not have and also usually has some simple hairs. The calyx-lobes are imbricate in M.

Flora Neotropica

venosa but this is sometimes hardto see. The closed bud of M. venosa is somewhat pointed while in M. myrcioides it is an indented hemisphere because the calyx-lobes curl strongly inwards. 31b. Myrceugeniamyrcioidesvar. acrophylla(Berg) Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 9. 1953. Figs. 8G-H, 19. Eugenia acrophvlla Berg, in Mart. Fl. Bras. 14(1): 575. 1859. Eugenia estrellensis Berg, in Mart. Fl. Bras. 14(1): 250. 1857. Type. Riedel s.n., "Habitat in nemoribus montium Serra d'Estrella in prov. Rio de Janeiro" (holotype, B, apparently lost; possible isotype, P). Luma estrellensis (Berg) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 529. 1941. Luma acrophvlla (Berg) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 531. 1941. ?Luma ulei Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 531. 1941. Type. Ule 1433, "Prov. Santa Catarina. Strauch in sumpfiger Capoeira bei Tubarao" (holotype, B, n.v.). Mvrceugenia estrellensis (Berg) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 9. 1953. ?Mvrceugenia mvrcioides var. ulei (Burret) Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 9. 1953. Myrceugenia mvrcioides var. paranensis Legrand, Darwiniana 11(2): 341. 1957. Type. Imaguire s.n. (Hatschbach 1826), Parana, Morretes (holotype, MVM, n.v.; isotypes, MBM, MICH). Mxrceugenia acrophvlla (Berg) Legrand, in Legrand et Klein, Flora Ilustr. Catar. [Mirt]: 342. 1970. Mvrceugenia ferreira-limana Klein et Legrand, in Legrand et Klein, Flora Ilustr. Catar. [Mirt]: 374. 1970. Type. Santa Catarina, Florian6polis, Morro Costa da Lagoa, Ilha de S. Catarina, 490 m, 31 Jan 1969, Klein & Souza Sob. 8079 (holotype, MVM). ?Mvrceugenia acrophvlla var. ulei (Burret) Legrand, in Legrand et Klein, Flora Ilustr. Catar. [Mirt]: 347. 1970.

Hairs greyish-white,reddish-brownor yellowish-brown,often all less than 0.3 mm long; leaves 4-16 cm long, 1.5-7 cm wide; blades membranousto subcoriaceous; bracteoles 2-8 mm long, 1-4 mm wide, 2-5.5 times as long as wide, densely pubescent without, densely pubescent to sparsely pubescent within, persistent or caducous before the fruit matures; calyx-lobes ovate to triangular,46.5 mm long, 3.5-6 mm wide, valvate or overlappingalong the narrowmembranous marginsof two lobes in the bud, otherwise coriaceous, sparsely pubescent distally to entirely glabrous within, densely pubescent to glabrous without; stamens ca. 250-500. Type. Riedel s.n., "Habitatad marginessilvarummontis Serrad'Estrellaprov. Rio de Janeiro, florebatJanuario-Februario"(holotype, LE, n.v.). Specimens examined. BRAZIL. Minas Gerais: Caldas, 1842, Regnell Ser. I no. 130 (US). Rio de Janeiro: Tingua, 5 Feb 1972, Emygdio 3085 (R); Jacarepagua, Represa do C-[?], 24 Jul 1933, Kuhlmann s.n. (RB); Serra da Estrella, without date, Saldanha 5233 (RB); Terez6polis, Parque Nacional Serra dos Orgaos, 18 Jun 1943, Dionisio & Otario 221 (RB), 22 Oct 1942, Duarte de Barros 1069 (RB); Petr6polis, 24 Feb 1915, Diogo 414 (R). Sao Paulo: Alto da Serra, Estacao Biologica, 29 Sep 1942, Kuhlmann s.n. (SP); Juquia, margem do Rio Assingui, 25 Jun 1941, Kuhlmann & Kuehn s.n. (SP). Parana: without specific locality, 23 Jan 1904, Dusen 3444 (R, US); Mun. Antonina, Rio Cotia, 17 Sep 1965, Hatschbach 12810 (F, HB, MBM, MVM, US); Mun. Antonina, Sapitanduva, 50 m, 31 Jan 1968, Hatschbach 18523 (MBM, MO, MVM, NY), 21 Mar 1972, Hatschbach 29330 (MBM, NY, UC); Mun. Campina Grande do Sul, Sitio do Belizario, 9 Aug 1967, Hatschbach 16803 (CTES, MBM, MVM), 1100 m, 16 Nov 1967, Hatschbach 17863 (MBM, MVM, UC); Mun. Campinas do Sul, Serra do Capivari, 2 Dec 1962, Hatschbach 9530 (H, HB, MBM, MICH, MVM); Mun. Guaratuba, Pedra Branca de Araraquara, 10 Mar 1963, Hatschbach 9772 (HB, MBM, MICH, MVM); Mun. Guaratuba, Morro do Morretes, 15 Feb 1964, Hatschbach 10972 (F, MBM, MICH, MVM); Mun. Guaratuba, Morro das Caieiras, 30-50 m, 23 Mar 1964, Hatschbach 11136 (MBM, MVM); Mun. Morretes, Rio Mae Catira, 24 Jan 1951, Hatschbach 7657 (HB, MBM, MICH, MVM, RB); Mun. Morretes, Estrada Graciosa, Grota Funda, 23 Feb 1947, Hatschbach 628 (MBM, MICH, MVM), 11 Apr 1948, Hatschbach 909 (MBM, MICH, MVM, PACA), 15 Feb 1964, Hatschbach 10972 (F, MBM, MICH, MVM), 6 Apr 1973, Hatschbach 31810 (C, MBM, NY, UC). 600 m, 13 Dec 1977, Landrum

Myrceugenia

2866, 2875, 2883 (MBM, MICH);Mun. Morretes,EstradaGraciosa,ca. 100m, 13Dec 1977,Landrum 2857, 2858, 2859, 2860, 2861, 2862 (MBM, MICH);Mun. Morretes,EstradaGraciosa,Boa Vista, 26 Dec 1962, Hatschbach 9832 (HB, MBM, MICH, MVM);Mun. Paranagua,Morrodo Tabaquara,27 July 1966, Hatschbach 14546(HB, MBM, MICH, MO, MVM, NY, US); Mun. Paranagua,Caioba, Morro do Boi, 70 m, 9 Mar 1964, Hatschbach 11119 (MBM, MVM); Mun. Paranagua,Praia do Mendanha,20-30 m, 1 Mar 1960, Hatschbach6785 (MBM, MVM);Mun. Pien, Trigolandia,9 Mar 1967, Hatschbach 16116 (MBM, MICH); Mun. Quatro Barras, Cerne, 23 Feb 1964, Hatschbach 10989(MBM, MVM);Mun. Sao Jose dos Pinhaes, Col. S. Andrade,2 Aug 1966, Hatschbach 14602 (C, CTES, MBM, MICH, MVM).SantaCatarina:Blumenau,Feb 1888, Ule 706 (US), Sep 1888, Ule 918 (US); Bom Retiro, Riosinho, 1000m, 23 Dec 1948, Reitz 2744 (H, MVM, NY, PACA, UC), 25 Nov 1956, Smith, Reitz & Klein 7932 (HBR, MVM, R, US); Mun. Brusque, Matado Hoffmann,30 Jun 1950, Veloso 16b (RB, US), 40 m, 18 Aug 1953, Klein567 (H, HBR, MVM, NY, UC, US); Mun. Chapeco, SeminarioDiocesiano, west of Chape9o,ca. 27?06'S,52?37'W,350-450 m, Smith & Klein 14035 (GH, HBR, MICH, MVM, NY, UC, US); Mun. Dionisio Cerqueira,13 km west of Rio Capetingabetween CampoEre and Dionisio Cerqueira,900-1000 m, 22 Feb 1957, Smith & Klein 11678 (HBR, MICH, MVM, NY, US, R); Mun. Ibirama,Estacao Florestal I. N. P., 300 m, 7 Mar 1956, Klein 1934 (H, HB, MBM, MVM, NY, UC, US); Mun. Ibirama,Horto FlorestalI. N. P., 350 m, 16 Aug 1956, Reitz & Klein 3595 (GH, H, MVM, NY, UC), 250 m, 4 Feb 1956, Reitz & Klein 2581 (H,

MVM); Mun. Irani, 1000 m, 28 Dec 1963, Reitz & Klein 16491 (HBR, H, MVM); Mun. Irani, Rio Irani, 900 m, 27 Feb 1964, Klein 4807 (H); Mun. Itajai, Morroda Ressacada, 150 m, 12 Aug 1955, Klein 1518 (H, MVM, US); Mun. Itajai, Morroda Fazenda, 50 m, 18 Feb 1954, Reitz & Klein 1551 (H, PACA, US); Mun. Itajai, Luis Alves, Braco Joaquim,300 m, 17 Jul 1954, Reitz & Klein 2400 (H, MICH, MVM, US); Mun. Itajai, Cunhas, 15 m, 10 Mar 1955, Klein 1187 (H, MVM, US), Klein 1196(H, MVM), 26 Jul 1955, Klein 1482 (H, HB, MBM, MVM);Mun. Joaqaba,15 km east of Ponte Serrada,700-900 m, 3 Jan 1957, Smith & Reitz 9873 (HBR, MVM, R, US); Mun. Joinvile, Estrada Dona Francisca,650 m, 21 Jun 1957,Reitz & Klein4451 (H, HB), 550 m, 22 Jan 1958,Reitz & Klein 6280 (H, MBM, NY, UC); Mun. Papanduva,north of the SerraGeral on the Estradade Rodagem, elev. 700-900 m, 6-7 Dec 1956, Smith & Klein8393 (HBR, MVM, R, UC, US); Mun. P6rto Uniao, 750 m, 25 Feb 1962,Reitz & Klein 12430(HBR, H, MVM, US), by new airporteast of P6rto Uniao, 750 m, 19 Dec 1956, Smith & Reitz 8834 (HBR, MICH, MVM, R); Mun. Santa Cecilia, a 30 km de Santa Cecilia, 20 Oct 1961, Pereira 6243 (HB, R, RB); Mun. Sao Jose, Serrada Boa Vista, 1200m, 4 Feb 1953, Reitz 5483 (H, HBR, MBM, MVM, NY, PACA, UC, US); Mun. Sao Franciscodo Sul, perto das Canoas [?], 21 Feb 1952, Reitz 4398 (H, MVM); Mun. Sao Francisco do Sul, Morrodo Campo Alegre, 1200 m, 20 Jan 1961, Reitz & Klein 10673 (HBR, H, MVM, NY, UC, US); Mun. Vidal Ramos, Sabia, 750 m, 7 Mar 1958, Reitz & Klein 6586 (H, MVM, NY, US). Rio Grandedo Sul: Sao Franciscode Paula, 9 Feb 1948,A. Mattos & Labouriaus.n. (RB); Mun. Rolante, Fazenda Englertnear Santa Tereza, ca. 1000m, 28 Dec 1977, Landrum3001 (MICH, PACA), withoutdate, Rainbo 56364 (MICH, MVM, PACA).

Mvrceugenia myrcioides var. acrophylla could only be confused with M. cu-

cullata. The two are compareddirectly in the discussion of that species. No type specimen has been seen for Eugenia acrophylla Berg, but that name has been applied to this variety by Legrand since 1953 and is retained here, at least until a type can be seen. If the type does belong to this entity, which seems likely given the description,then E. acrophylla is the name which should be used as a basionym. If it does not belong here then two other epithets will have to be considered: ulei for which no type has been found but which is earlier; and paranensis for which the type has been found and which definitely belongs to this variety. 32. Myrceugeniapilotantha (Kiaerskou) Landrum, Brittonia 32(3): 374. 1980. Eugenia pilotantha Kiaerskou, Enum. Myrt. bras.: 169. 1893.

Small tree or shrub 1.5-12 m high; hairs reddish-brown,usually a mixture of simple and asymmetrically dibrachiate; twigs densely pubescent when young, glabrescentwith age, the barkbrown or light grey, rarelybecoming flaky; leaves elliptic, lanceolate, ovate, obovate or oblanceolate, 2-9.5 cm long, 0.7-3.6 cm wide, 1.8-4 times as long as wide, the blade sparselypubescent to glabrous(rarely densely pubescent) above, densely pubescent when young, becomingless so with

Flora Neotropica

Table VI Comparisonof Legrand's Division of the Myrceugeniamiersiana Complex with That Presented Here Landrum M. miersiana (Gardner) Legrand et Kausel

M. pilotantha (Kiaerskou) Landrum var. pilotantha

M. pilotantha var. major

Legrand M. miersiana (Gardner) Legrand et Kausel var. miersiana M. fuscovelutina (Burret) Legrand et Kausel M. nothorufa var. venosa Legrand M. miersiana var. venosa (Berg) Legrand et Klein, as to name only, probably not including the type of E. miersiana [var.] venosa M. miersiana var. macahensis (Burret) Legrand M. ramboi Legrand M. nothorufa Legrand var. nothorufa M. nothorufa var. major Legrand

age below, the midvein densely pubescent above and below, becoming less so with age, the hairs of the lower surfaceusually somewhat appressed;apex acute, acuminate,or rounded,the midveinoften slightlyexcurrent;base acute, cuneate, obtuse, or acuminate;petiole subterete or only slightly sulcate above, densely pubescent, 2-8 mm long, 0.5-1 mm thick; midvein impressedfor entire length or only distally above, prominentbelow; lateral veins indistinct or up to about 10 pairs clearly visible; marginal veins equallinglaterals in prominenceor less distinct; blades coriaceous to subcoriaceous, olive-green to light grey-green, often tinged with reddish-brownabove, about the same color but lighter below; peduncles uniflorous, 0-17 mm long, 0.5-1 mm wide, densely pubescent, solitary or up to 3 in a row in the leaf axils; bracteoles lanceolate to linear, 1.5-5 mm long, 0.5-2 mm wide, (1.6-)2-4.8 times as long as wide, densely pubescent within and without, subcoriaceous, claspingthe hypanthium;calyx-lobes ovate, oblongovate to triangular, 1.6-4.5 mm long, 1-3.5 mm wide, 1-1.7 times as long as wide, densely pubescent without, densely pubescent within except sometimes at base; hypanthiumobconic, densely pubescent, 1-3 mm long; disk ca. 1-3 mm across, sparsely to densely pubescent; style 4-7 mm long, glabrous to sparsely pubescent; stamens ca. 100-230, 3-8 mm long; anthers 0.3-0.6 mm long when dry; petals suborbicular,densely pubescent without, 2-4 mm in diam.; ovary 24-locular;ovules 4-15 per locule; fruit darkpurple, 5-10 mm in diam.; seeds ca. 3-6 per fruit. Myrceugenia pilotantha appears to be most closely related to M. miersiana.

The two species probablycomprise a close-knit phylogenetic unit definedby the combinationof pubescent petals, simple or a mixture of simple and dibrachiate hairs, and petioles that are terete or scarcely channeled. Withinthe group I have chosen to recognize two species: M. pilotantha with two varieties and M. miersiana with one. Legrand(1953, 1957, 1961)and Legrand& Klein (1970) divided the group in a differentfashion which is comparedwith mine in Table VI. The three entities recognized here form a progression with Myrceugeniapilotantha var. major and M. miersiana the extremes, and M. pilotantha var. pil-

otantha intermediatebetween them. The extremes grow sympatrically,are quite distinct and hybridize rarely if at all. Myrceugenia pilotantha var. pilotantha is

for the most part separatedgeographicallyfrom both the others and only seems

Myrceugenia

to merge morphologicallywith M. pilotantha var. major in the vicinity of Sao Paulo. It most closely resembles M. miersiana in the state of Rio de Janeirobut so far has been separable from that species. The two species are compared directly with each other in Key K. Key to Varieties of Myrceugenia pilotantha 1. Calyx-lobesblunt; leaves elliptic to lanceolate, usually 4-8 cm long, usually 3-4 times as long as wide; floweringmainlyfrom Februaryto May;fruits maturingmainlyfrom July to September;growingmainlybelow 900 m; Santa Catarinato Rio de Janeiro. a. var. pilotantha. 1. Calyx-lobesacute; leaves elliptic to obovate or oblanceolate,usually 2-5 cm long, usually 2-3 times as long as wide; floweringmainlyfrom December to February;fruits maturing mainlyfrom Septemberto November;growingmainlyabove 900 m; Paranato Rio Grande b. var. major. do Sul.

32a. Myrceugenia pilotantha var. pilotantha

Fig. 20.

Eugeniapilotantha Kiaerskou,Enum. Myrt. bras.: 169, as to type, 1893. Lumapilotantha (Kiaerskou)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 527. 1941. Luma macahensis Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 527. 1941. Type. Glaziou 18249,

"Brasilien:Alto Macah6de Nova Friburgo,Staat Rio de Janeiro"(holotype, B, n.v., isotype, C). Myrceugeniamiersiana var. macahensis (Burret)Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo 2(28):8. 1953.

Leaves elliptic, lanceolate or ovate, 2-9.5 cm long, 0.7-3.6 cm wide, 1.8-4 times as long as wide; peduncles 1-17 mm long, 0.5-1 mm wide; bracteoles lanceolate to linear, 1.5-4 mm long, 0.5-1 mm wide, (1.6-)2-4.8 times as long as wide; calyx-lobes blunt, 1.6-3.5 mm long, 1-2.8 mm wide, 1-1.7 times as long as wide. Type. Glaziou 16052(holotype, C; Field Museumneg. 20971of holotype, photo at MICH). Specimensexamined.BRAZIL.Rio de Janeiro:Petr6polis,Quitandinha,1948,Goes & Octavio 10 (RB); Petr6polis, Rocio mais ou menos, 16 Mar 1968, Sucre 2462 (MICH, MO, RB); Petr6polis, Cascade d'Itamaraty,20 Feb 1873, Glaziou6543 (C, US); estradapara o Pico de Tijuca, 550 m, 31 Mar 1971, Sucre 7567 (MICH);Tijuca, 2 Apr 1868, Glaziou491 (US); Tijuca,pres de la Cascada,2 Apr 1868, Glaziou2591 (C); Bico do Papagaio,13 Mar 1972, Laroche & Almeida 1365(MICH,RB); ParqueNacionalTijuca,Bico do Papagaio(ca. 22?50'S,43?20'W),800-1000m, 24 Oct 1977,Landrum 2216 (MICH, RB); Alto da Boa Vista, entre Mesa do Imperadore Alto da Boa Vista, 24 Mar 1959, Duarte 4649 (HB, MICH, RB); ParqueNacional Tijuca, Mesa do Imperador(ca. 22?50'S,43?20'W), 400-650 m, 12 Oct 1977, Landrum2021, 2023 (MICH, RB); estrada do Redentador, 14 Apr 1941, Kuhlmanns.n. (MICH, MO, RB): Sumar6,Estradado Corcovado, 530 m, 23 Feb 1972, Almeida 1265(MICH,MO, RB); withoutspecificlocality, Glaziou1103(C, MO), Glaziou8712 (C). Sio Paulo: Alto da Serra, Estacao Biol6gica, 10 Jan 1919, Hoehne s.n. (SP), 7 Jan 1919, Gehrts.n. (SP); Alto da Serra, Via Anchieta, pr6ximo do Posto de Pedagio, 30 Oct 1961,J. Mattos s.n. (SP), ca. 800 m, 10 Dec 1977, Landrum2849 (MICH, SP); Esta9ao de Campo Grande(Estrada de Ferro SantosJundiai),Estatao Biologica, 6 Dec 1960, J. Mattos 10568, 10569 (SP). These specimens from Sao Paulo are more or less transitionalbetween the two varieties of Myrceugeniapilotantha. Santa Catarina:Mun. Rancho Queimado,Serra da Boa Vista, 700 m, 13 Apr 1963, Reitz & Klein 10992 (HBR, MVM, NY, UC, US), 12 Aug 1960,Reitz & Klein 9733 (H, HBR, MVM);Mun. Florian6polis, Morro Costa da Lagoa, 490 m, 23 May 1968, Klein & Bresolin 7717 (MVM); Mun. Florian6polis, MorroRio Vermelho,250 m, 27 Jun 1968, Klein & Bresolin 7771 (MVM), 300 m, 6 Aug 1968, Klein 7830 (MVM).

Myrceugenia pilotantha var. pilotantha is a small tree to ca. 12 m high, typi-

cally growingin warmhumidforests from ca. 250 to 900 m nearthe Atlanticcoast. A map of its distributionis shown in Fig. 20. It flowers mainlyfrom Februaryto May and the fruits probablymaturefrom July to September. Aside from Myrceugenia miersiana this variety might be confused with M. rufescens, which differs in having its petioles channeled and its petals and the

Flora Neotropica 50

. .:

.- ... .'

. :

?.3 .

. .:

... . ..

.... .

pi;

ot.nh !.

var..

FIG. 20.

Distribution of Myrceugenia pilotantha.

lower surfaces of mature leaves sparsely pubescent to glabrous. In addition M. rufescens flowers mainly in September and October and normally has some bracteate shoot inflorescences. 32b. Myrceugenia pilotantha var. major (Legrand) Landrum, Brittonia 32(3): 374. 1980. Figs. 81, 20. Mvrceugenia nothorufa var. major Legrand, in Legrand et Klein, Flora Ilustr. Catarinense [Mirt]: 414. 1970. Myrceugenia ramboi Legrand, Darwiniana 11: 320. 1957. Type. Rambo 53920, Rio Grande do Sul, Sao Francisco de Paula, Taimbesinho (holotype, MVM; isotype, PACA). Myrceugenia nothorufa Legrand, in Legrand et Klein, Flora Ilustr. Catarinense [Mirt]: 409. 1970. Type. Reitz & Klein 6030, Santa Catarina, Campo Alegre, Morro do Iquererim, 1200 m, 9 Jan 1958 (holotype, MVM; isotype, US).

Leaves elliptic, obovate or oblanceolate, 2-6.5 cm long, 0.8-2.8 m wide, 1.83(-4) times as long as wide; peduncles 0-5 mm long, ca. 1 mm wide; bracteoles lanceolate, 2-5 mm long, 0.7-2 mm wide, 2.5-3.6 times as long as wide; calyxlobes sharply acute, 2.5-4.5 mm long, 2-3.5 mm wide, 1-1.5 times as long as wide. Type. Reitz & Klein 8403, Santa Catarina, Bom Jardim, Curral Falso, 1500 m, 19 Feb 1959 (holotype, MVM, isotype, HBR). Specimens examined. BRAZIL. Parana: Mun. Quatro Barras, Rio do Corvo, 900 m, 14 Dec 1969, Hatschbach 23240 (C, MBM, MVM, UC); Mun. Campina Grande do Sul, Alto da Serra, 19 Nov 1959, Hatschbach 6531 (MBM, MVM); Mun. Sao Jos6 dos Pinhais, road to Guaricana (ca. 25?40'S, 49?W), 900-1000 m, 4 Nov 1977, Landrum 2370 (MBM, MICH), 15 Dec 1977, Landrum 2910 (MBM, MICH). Santa Catarina: Mun. Campo Alegre, Morro do Iquererim, 1500 m, 5 Sep 1957, Reitz & Klein 4769 (HBR, MVM, NY, US), 1300 m, 10 Jan 1958, Reitz & Klein 6110 (HBR, MVM, US); Mun. Blumenau, Morro Spitzkopf, 900 m, 6 Feb 1960, Reitz & Klein 9544 (HBR, MVM, NY, UC, US); Mun. Rancho Queimado, Serra da Boa Vista, 1200 m, 24 Oct 1957, Reitz & Klein 5431 (HBR, MVM, US), 1000 m, 25 Jan 1961, Reitz & Klein 10775 (HBR, MVM, US); Mun. Bom Retiro, Campo dos Padres, 1300-1400 m, 22 Nov 1956, Smith & Klein 7863 (US), 1650 m, 17-19 Nov 1956, Smith Reitz & Klein 7729 (F, HBR, MVM, R, RB, UC, US), 25 Jan 1957, Smith & Reitz 10466 (HBR, MICH, MVM, R, US); Mun. Lajes, between the crest of the Serra Geral and Encruzilhada, 900-1000 m, 2 Dec 1956, Smith & Klein 8022 (HBR, MVM, R, UC); Mun. Lajes, 2-11 km past crest of Serra

Myrceugenia

Geral on road to Otacilio Costa from Rio do Sul, ca. 1000 m, 21 Nov 1977, Landrum2633, 2649, 2650, 2651, 2652 (MICH, MBM);Mun. LauroMuller,VargemGrande,400 m, 17 Dec 1958,Reitz & Klein 8093 (HBR, MVM, UC, US). Rio Grandedo Sul: Mun. Cambara,Fortaleza(ca. 29?S,50?W), ca. 1100m, 27 Dec 1977, Landrum2972, 2976 (MICH, PACA).

This variety is a shrub or small tree up to ca. 6 m high, typically growing in Araucaria forests and along the eastern edge of the planalto at elevations of 9001650 m. A map of its distribution is shown in Fig. 20. It flowers mainly from December to February and the fruits mature from September to November. Myrceugenia pilotantha var. major is rather similar to M. acutiflora and is probably a close relative. The two are compared in the discussion of M. acutiflora. 33. Myrceugenia miersiana (Gardner) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 8. 1953. Fig. 21. Eugenia miersianaGardner,LondonJ. Bot. 4: 103. 1845. Eugenia miersiana[var.]a costata Berg, in Mart.Fl. Bras. 14(1):574. 1859.(Inadmissiblename to be replacedby E. miersiana var. miersiana because it is based on the same type as E. miersiana Gardner.) ?Eugenia miersiana [var.] ,3 venosa Berg, in Mart. Fl. Bras. 14(1): 574. 1859. Type. Riedel s.n.,

"Habitatin prov. Rio de Janeiro," "v. in hb. hort. Petrop." (holotype, LE, n.v.). ?Eugeniamiersiana [var.] y glomerata Berg, in Mart. Fl. Bras. 14(1):574. 1859.Type. Riedel s.n., "Habitatin prov. Rio de Janeiro," "v. in hb. hort. Petrop." (holotype, LE, n.v.). ?Eugeniamiersiana[var.] 5 membranaceaBerg, in Mart.Fl. Bras. 14(1):574. 1859.Type. Riedel s.n., "Habitatin prov. Rio de Janeiro," "v. in hb. hort. Petrop." (holotype, LE, n.v.). Luma miersiana(Gardner)Burret,Notizbl. Bot. Gart. Berlin-Dahlem15: 527. 1941. LumafuscovelutinaBurret,Repert. Spec. Nov. Regni Veg. 50: 51. 1941.Type. Tessmann6115, "Sud-Brasilien:Nord-Parana,Gebiet des mittlerenIvahy, Regenwald" (holotype, B, apparentlylost; apparentisotype, RB, labeled Tessmann115 but otherwise correspondingto protologue). Myrceugeniafuscovelutina (Burret)Legrandet Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 13. 1953. Myrceugenianothorufa var. venosa Legrand, in Legrandet Klein, Flora Ilustr. Catarinense [Mirt]:413. 1970.Type. Reitz 5476, SantaCatarina,RanchoQueimado,Serrada Boa Vista, 1300m, 4 Feb 1953(holotype, MVM;isotypes, HBR, US). ?Mvrceugeniamiersiana var. venosa (Berg) Legrandet Klein, Flora Ilustr. Catarinense[Mirt]: 399. 1970.

Shrub or tree 2-10 m high; hairs golden-brown to rusty-brown or yellowish, all simple or nearly so; twigs densely pubescent when young, the hairs persisting until the first bark falls, the bark of older twigs light grey to whitish; leaves narrowly to broadly elliptic, less often tending to be ovate, lanceolate or obovate, 2.5-10 cm long, 1-3 cm wide, 2-4.5 times as long as wide, the upper surface glabrous or nearly so on the blade, densely pubescent along the midvein, the lower surface pubescent when young, sometimes glabrescent with age, the hairs of the lower surface mainly erect; apex acute or less often rounded or acuminate, the midvein often excurrent; base acute; petiole subterete or only slightly sulcate, densely pubescent, 2-7 mm long, 1-1.5 mm thick; midvein impressed above, prominent below; lateral veins prominent, up to about 15 pairs clearly visible; marginal veins about equalling laterals in prominence and arcing between them; blades coriaceous to subcoriaceous, light grey-green to olive-green, often tinged with reddish-brown above, about the same color and somewhat lighter below; peduncles uniflorous, 1-15 mm long, 0.5-1 mm wide, densely pubescent, solitary or up to 3 in a row in the leaf axils; bracteoles ovate to lanceolate, 1.5-6 mm long, 0.7-1.2 mm wide, 1.7-5 times as long as wide, densely pubescent within and without, clasping the hypanthium; calyx-lobes ovate to triangular, blunt or less often acute, 2-3.6 mm long, 2.2-3.8 mm wide, 0.7-1.2(-1.4) times as long as

Flora Neotropica

100

wide, densely pubescent without, densely pubescent within except sometimes at the base; hypanthium obconic, densely pubescent, 1.5-3 mm long; disk 2-3.8 mm across, moderately to sparsely pubescent; style 5-9 mm long, sparsely pubescent to glabrous; stamens 100-320, 3-9 mm long; anthers 0.4-0.7 mm long when dry; petals more or less orbicular, 2-4 mm in diam., densely to sparsely pubescent without, the margin sometimes irregular; ovary 3-4-locular; ovules 716(-22) per locule; fruit globose, 6-12 mm in diam., dark purple, sparsely pubescent; seeds 1-5 per fruit, round to oblong, 3-6 mm long. Type. Gardner 5712, "Woods in the Organ Mountains, at an elevation of about 3000 ft. Fl. March" (holotype, K). Specimens examined. BRAZIL. Minas Gerais: Fazenda da Gramma,900 m, 28 Jan 1930, Mexia 4257 (MO, RB); Camanducaia,SerrariaEsperanfa, 30 Jul 1949,Kuhlmanns.n. (SP). Rio de Janeiro: ParqueNacionalItatiaia,Rio CampoBelo (ca. 22?30'S,44?35'W),ca. 1000m, 17 Oct 1977, Landrum 2086 (MICH,RB); Petr6polis,Estradado Contorno,ca. 700 m, 23 Mar 1968,Sucre2539 (HB, MICH, RB). Sao Paulo: Camposdo Jordao,Reserva Florestalde Camposdo Jordao(ca. 22?45'S,45?30'W), ca. 1600m, 5 Dec 1977, Landrum2823 (MICH, SP); Camposdo Jordao, ParqueEstadual,23 Apr 1974,J. Mattos 15776(SP); Sao Luiz de Paraitinga,rodoviaUbatuba-Taubate,2 May 1961,J. Mattos 8945 (MBM, SP). Parana:Mun. CampinaGrandedo Sul, Serrada Lapinha,5 May 1963,Hatschbach 9977 (F, HB, MBM, MICH, MVM), Hatschbach 10035(F, HB, MICH);Mun. CampinaGrandedo Sul, Caminhoao Cerro Verde, 1100m, 21 May 1967, Hatschbach 16467 (MICH);Mun. Piraquara, Banhados, 11 Nov 1943, Hatschbach 103 (MBM, MICH, MVM); Mun. Piraquara,Novo Tirol, 26 Apr 1964, Hatschbach 11244(F, HB, LP, MBM, MVM);Mun. Piraquara,Mananciasda Serra(ca. 25?30'S,49?W),ca. 1000m, 31 Oct 1977,Landrum2235 (MBM, MICH);Mun. Sao Jose dos Pinhais, Col. S. Andrade, 12 Aug 1966, Hatschbach 14593 (MBM, MICH, MVM); Mun. Sao Jose dos Pinnhais, road to Guaricana,900-1000 m, 4 Nov 1977, Landrum2404, 2408 (MBM, MICH);Mun. Mandirituba,29 Jun 1948,Hatschbach975 (MBM, MICH,MVM,PACA);Mun. Guaratuba,Garuva, 24 Mar 1957, Hatschbach 4019 (HBR, MBM, MICH, MVM, PACA); Mun. Sao Joao do Triunfo, Fazenda Sao Joao da Fiat Lux Cia, ca. 840 m, 21 Jul 1966, Lindeman& de Hass 1892 (MBM, NY, RB); Mun. Gal. Carneiro,Cab. Rio Iratim,18 Oct 1966,Hatschbach 14997(C, MBM, MICH,MVM, UC); Mun. Rio Negro, Pien, 24 Apr 1959, Hatschbach5634 (HBR, MBM, MVM);Capao Grande, 30 Apr 1909, Dusen 8065 (MICH); Fazenda Experimentalda Escola de Agronomia, 16 Jul 1968, Imaguires.n. (CTES). SantaCatarina:Mun.Rio do Sul, Matador,350 m, 17Apr 1959,Reitz & Klein 8760 (H, HBR, MVM, NY, UC, US); Mun. Vidal Ramos, Sabia, 750 m, 7 Apr 1958, Reitz & Klein 6645 (H, HB, HBR, MBM, MVM, NY, UC, US). Rio Grandedo Sul: Sao Franciscode Paula,8 Feb 1948, A. Mattos & Labouriaus.n. (MVM, RB); Mun. Sio Francisco de Paula, 5 km along road to P6rto Alegre (ca. 29?30'S, 50?15'W), ca. 1000 m, 26 Dec 1977, Landrum 2942, 2946, 2949 (MICH,

PACA);Mun. Cambara,Fortaleza(ca. 29?S,50?W),ca. 1100m, 27 Dec 1977,Landrum2978 (MICH, PACA).

Myrceugenia miersiana can easily be confused with M. venosa; in fact Legrand described M. miersiana var. lanceolata which I believe to be a synonym of M. venosa. Myrceugenia miersiana is compared directly with M. venosa and M. pilotantha, another apparently closely related species, in Key K. Myrceugenia miersiana flowers mainly from February to April and the fruits probably mature from August to October. It seems to grow in Araucaria forests and along the eastern edge of the planalto. A map of its distribution is shown in Fig. 21. 34. Myrceugenia myrtoides Berg, Linnaea 27: 133. 1856 (name only); in Mart. Fl. Bras. 14(1): 211. 1857. Figs. 21, 33C. Myrceugenia mvrtoides [var.] / stricta Berg, in Mart. Fl. Bras. 14(1): 211. 1857. Type. Sellow

s.n., "in fruticetis ripariisad Rio Pardo in Montevideo, floret Novembri" (holotype, B, apparentlylost; isotype, W, hereby designatedas lectotype; isolectotype, P). (Because the type of this taxon has been chosen as the lectotype of M. myrtoides, the name would be replaced by M. myrtoides var. myrtoides.)

Myrceugeniamyrtoides[var.]a conferta Berg, in Mart.Fl. Bras. 14(1):211. 1857.Type. Sellow s.n., "ad ripas. fluminisYrapua"(originalspecimenB, apparentlylost; isotype, W, hereby designatedas lectotype); Sellow s.n., "in silvis in Montevideo" (paratype,B, apparently

101

Myrceugenia 50 20

:-'

FIG. 21.

Distributions of Myrceugenia miersiana and M. myrtoides.

Myrceugenia montevideensis Berg, innaea 27: 133. 1856 (name only); in Mart. Fl. Bras. 14(1):!' of isotype at P, photo at MICH). ?Myrceugenia montevideensis Berg, Linnaea 27: 133. 1856 (name only); in Mart. Fl. Bras. 14(1):

ofsimple and asymmetricallyBurret, and Veg. Spe twisted .9Luma cinnamomeotomentosa dibrachiate, Repert. . ?Regni 50: 51.together; 1941. Type. twigs Artangled chavaletaMvrt.n. 17, "Uruguay:MeloGueso"(holotype,B, apparently lost).

of simpleandasymmetrically dibrachiate, twistedandtangledtogether;twigs

102

Flora Neotropica

densely lanate when young, glabrescentwith age, the bark smooth, whitish-grey; leaves densely lanate below, puberulentabove, losing most or all hairs within a year, ovate, lanceolate, elliptic to suborbicular(0.8-)1-2.5 cm wide, 2-6 cm long, 1.4-4.5 times as long as wide, the margin often slightly revolute; apex acute (rarelyrounded),the midveinusually slightlyexcurrent;base roundedto cuneate; petiole channeled, 2-4 mm long, 0.5-1.5 mm thick; midvein only slightly impressed above, more puberulentthan the surroundingblade, prominentbelow; lateral veins indistinctor up to ca. 20 pairs faintly visible; marginalveins equalling or less distinct than the laterals; blades dull yellow-green to grey-green above, lighter to almost white beneath, coriaceous to chartaceous; peduncles uniflorous(rarely bearinga three flowered dichasium),flattened, 2-15 mm long, 1-1.5 mm wide, densely lanate, solitary or in pairs in the leaf axils; bracteoles lanceolate, 4.5-8 mm long, 1.6-3 mm wide, 2.5-4 times as long as wide, chartaceous, densely lanate within and without, claspingthe hypanthium;calyx-lobes deltoid to lanceolate, 4-7 mm long, 3-5 mm wide, 1.4-2.5 times as long as wide, chartaceous, densely lanate within and without, concave proximallyor not at all concave; hypanthium obconic, densely lanate, 2.5-4 mm long; disk 3-5 mm across, densely to sparsely covered with hairs; style 9-14 mm long, sparsely pubescent proximally;stamens 120-250, 5-15 mm long; anthers 0.4-0.7 mm long when dry; petals suborbicular,concave, 3-6 mm in diam.; ovary 2-4-locular; ovules 9-14(-17) per locule; fruit light grey, 5-6 mm in diam. or perhapslarger. Type. Sellow s.n., "in fruticetis ripariis ad Rio Pardo in Montevideo, floret Novembri," type of Myrceugeniamyrtoides [var.] / stricta Berg, hereby designated as the lectotype of Myrceugeniamyrtoides Berg (lectotype, W; isolectotype, P). Specimensexamined.BRAZIL.Rio Grandedo Sul: Canoas,3 Jun 1949,Rambo41820 (H); Esteio, 24 Nov 1948, Rambo 38259 (H, LIL); Gravatai,Cachoerinha,7 Jan 1949, Rambo 39588 (H); Montenegro, Estaaio Pareci, 14 Jan 1949, Rambo 39714 (H, PACA);P6rto Alegre, Barrado Ribeiro, 5 Apr 1950, Rambo 46640; P6rto Alegre, Dec 1898, Reineck & Czermak 263 (H, MVM, W); P6rto

Alegre, zwischen Sao Joao und Navegantes, 15 Dec 1897, Reineck & Czermak163 (G, P); Sao Leopoldo, Steinkopf, 20 Dec 1949, Rambo 39039 (H); Sao Leopoldo, Rio dos Sinos, 10 Dec 1948, Rambo38694 (H, HBR);Sao Leopoldo, 10Oct 1946,Henz s.n. (MO, NY, SI); Sao Leopoldo,Arroio Manteiga,29 Dec 1977, Landrum3023, 3024, 3025 (MICH, PACA);Sao Leopoldo, Club de Pesca e Ca9a, 29 Dec 1977, Landrum3043, 3044, 3045, 3046 (MICH, PACA); Quarai,Jarau, 10 Jan 1945, Rambo26094 (F, MICH,MO, MVM,PACA, W); entre Pelotas e Sao Louren9o,4 Feb 1961,Pereira 6798 (HB, MVM, NY, RB); without specific locality, 1833, Gaudichaud1328 (P). URUGUAY. Tacuaremb6,Valle Eden, 11 Jan 1965, Del Puerto 3935 (US), 22 May 1964, Del Puerto & Marchesi3602 (MVM);CerroLargo,CerroGuazunzambu,Feb 1928,Schroders.n. (MVM).

The type of Myrceugeniamyrtoides [var.] 3 stricta Berg, has been chosen as the lectotype of M. myrtoides because Berg cited only one collection for this variety and thus it can be easily typified. For M. myrtoides [var.] a conferta Berg cited two Sellow collections. One collection from "fluminis Yrapua" is probablyfrom the Arroio Irapuain the Mun. Cachoeirado Sul, in Rio Grandedo Sul. This has been selected as the lectotype of M. myrtoides [var.] a conferta. The second collection cited by Berg is from "Montevideo," probably meaning somewhere in Uruguay. A specimen at G may be an isoparatype. This species is known only from Rio Grandedo Sul south of the planalto, and from northeasternUruguay where it seems to be a shrubgrowing along the ecotone between forests and campos. A map of its distributionis shown in Fig. 21. It is probably most closely related to Myrceugenia brevipedicellata, M. oxysepala

and M. leptocalyx but is easily distinguishedfrom all the other species of the genus by its long narrow calyx-lobes and abundantlylanate flowers, twigs and

Myrceugenia

103

lower leaf surfaces. It flowers from October to January and the fruits probably mature during the winter months from June to September. 35. Myrceugenia hoehnei (Burret) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 5. 1953. Fig. 22. Luma hoehnei Burret,Repert. Spec. Nov. Regni Veg. 50: 54. 1941.

Shrub 1-2.5 m high; hairs reddish-brown, simple and dibrachiate; twigs densely pubescent when young, glabrescent with age; leaves sparsely pubescent to puberulent above and below, glabrescent with age, elliptic to obovate, 2-4 cm long, 0.8-1.7 cm wide, 2.5-3.7 times as long as wide; apex acute, obtuse or acuminate, usually slightly mucronate; base acute, acuminate or cuneate; petiole channeled, pubescent, 1-3 mm long, 0.5-1 mm thick; midvein impressed above, prominent below; lateral veins indistinct or up to ca. 20 pairs distinguishable but not prominent; marginal veins equalling laterals in prominence; blades light to dark grey-green above, lighter grey-green to yellow-green below, coriaceous, the surfaces dull; peduncles uniflorous, flattened, 5-10 mm long, 0.5-1 mm wide, densely pubescent, solitary or in pairs in the leaf axils; bracteoles narrowly lanceolate to linear, ca. 2.8-6.2 mm long, 0.6-1 mm wide, 3.5-9 times as long as wide, subcoriaceous to coriaceous, densely pubescent without, less pubescent to glabrous within, loosely clasping the hypanthium; calyx-lobes acute, triangular to ovate, ca. 3-4.5 mm long, ca. 1.5-3 mm wide, 1.2-2 times as long as wide, densely pubescent without, pubescent distally to glabrous proximally within; hypanthium densely pubescent, ca. 1.5-2.5 mm long; disk ca. 2-2.5 mm across, sparsely pubescent; style 4-5 mm long; stamens ca. 100, 3-6 mm long; anthers 0.4-0.5 mm long when dry; petals ca. 1.5-2.5 mm in diam.; ovary 3-locular as far as known; ovules 5-10 per locule; fruit dark blackish-brown when dry, pubescent, 4.5-6.5 mm in diam.; seeds probably few per fruit, 3-4 mm in diam. Type. A. Gehrt s.n., Sao Paulo, Raiz de Serra (holotype, SP-17200). Specimensexamined.BRAZIL. Sao Paulo:Paranapiacaba,EstagaoBiol6gica, 18 Mar 1963, Han-

dro 1056 (SP), 5 Mar 1964, J. Mattos 11461 (MBM, MICH, SP, US), 3 Mar 1964, J. Mattos 11719

(C), 27 Jul 1967, Mattos 14838(SP), 26 Jul 1971,J. Mattos 15623(SP). Santa Catarina:Mun. Campo Alegre, lower slopes of Morro Iquererim, 1000-1300m, 9 Dec 1956, Smith & Klein 8494 (HBR, MVM, RB).

Legrand has used the name Myrceugeniafilibracteata (Burret) Legrand (1961) for this species, an epithet which is based on Luma filibracteata Burret (1941b). The holotype of Luma filibracteata, which was at B, has probably been lost and no isotype has been found. Although the description corresponds to M. hoehnei in some ways, the leaves are said to be 6-8 cm long and lanceolate or oblanceolate, whereas in M. hoehnei they are 2-4 cm long and mainly elliptic. Furthermore Burret indicates that L. filibracteata is similar to L. miersiana (=M. miersiana) and L. myrcioides (=M. myrcioides) and makes no mention of L. hoehnei which he describes two pages later. I think that L. filibracteata is most probably some form of M. miersiana, M. pilotantha or M. rufescens, or perhaps another species. If it turns out that it is a synonym of M. hoehnei, then one of the names will have to be chosen because the basionyms appear in the same publication. A good case could be made for using M. hoehnei because it has been in use in the genus for a longer time. Myrceugenia hoehnei might be confused with M. pilotantha but can be distinguished by its channeled petiole and petals that are not pubescent. From M. rufescens, another similar species, it can be distinguished by its calyx-lobes that

104

Flora Neotropica

become reflexed as they mature and that are not concave throughoutthe entire inner surface;its hairs that are a mixtureof simple and dibrachiate,not all simple as they usually are in M. rufescens; and its floweringperiod which is from March to July not August to October. As far as is known, Myrceugenia hoehnei inhabits the misty forests of the eastern edge of the planalto. A map of its distributionis shown in Fig. 22. Only one fruitingspecimen is known, that collected in December. 36. Myrceugeniabrevipedicellata(Burret) Legrand et Kausel, in Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 4. 1953. Figs. 22, 33D. Luma brevipedicellata Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15: 527. 1941.

Shrubor small tree 2-3 m high;pubescence reddish-brownto whitish, the hairs simple and dibrachiate,the dibrachiatehairs usually quite asymmetrical;twigs densely pubescent when young, glabrescent with age, the bark whitish-grey; leaves elliptic to narrowlyelliptic, 2.4-5 cm long, 0.7-1.5 cm wide, 2.6-4 times as long as wide, sparselyto moderatelypubescent below when young, glabrescent with age, sparsely pubescent to glabrousabove; apex acute to acuminate, often slightly mucronate;base cuneate; petiole channeled, densely to sparsely pubescent, 1-3 mm long, ca. 1 mm thick; midvein impressed above, prominentbelow; lateral veins indistinctor up to ca. 12 pairs, barely visible; marginal veins indistinct; blades darkor lightgrey-greenabove, lightergrey-greenor yellowish-green below, coriaceous; peduncles uniflorous,flattened,0-3 mm long, ca. 1 mm wide, densely pubescent, solitary or in pairs in the leaf axils; bracteoles lanceolate, 4.5-6.5 mm long, 1.5-2 mm wide, 2.6-3.7 times as long as wide, densely pubescent without, sparselypubescent within, subcoriaceous,claspingthe hypanthium, completely covering the young bud, the apex acuminate;calyx-lobes lanceolate, 2.5-4.5 mm long, 1-2 mm wide, ca. 2-3 times as long as wide, densely pubescent within and without, often concave proximally; hypanthiumdensely pubescent, ca. 2 mm long; disk ca. 2.5 mm across; stamens 130-190, 4-8 mm long; anthers ca. 0.3-0.5 mm long when dry; petals glabrousor nearly so, ca. 3 mm in diam.; ovary 2-3-locular; ovules 4-10 per locule; fruit unknown. Type. J. T. de Moura 777, "Staat S. Paulo, Campos do Jordao, 1800 m, am Gipfel" (holotype, presumably B, apparently lost; an isotype perhaps will be found in a Brazilianinstitution). Specimens examined. BRAZIL Sao Paulo: Campos do Jordao, 15 Dec 1952, Capell s.n. (RB); Campos do Jordao, Fazenda da Guarda (Reserva Florestal), 17 Dec 1966, J. Mattos 14736 (SP); Campos do Jordao, Reserva Florestal de Campos do Jordao (ca. 22?45'S, 45?30'W), ca. 1600 m, 5 Dec 1977, Landrum 2820, 2821, 2830 (MICH, SP).

Althougha type specimen has not been found for Luma brevipedicellata,Burret's description fits this species fairly well. Unless a type specimen is found which does not belong to this species, it is probablybest to follow Legrandand Kausel in using the epithet brevipedicellata. Myrceugeniabrevipedicellatais apparentlyan endemic of the Campos do Jordao region in Sao Paulo where it was found growing at the borders between wet campos and Araucaria forests. Its closest relatives seem to be M. oxysepala and M. leptocalyx, both more southern species. From these it can be easily distinguished by its narrowleaves and by various floralcharacters. Myrceugeniabrevipedicellata flowers in December and Januaryand probablyfruits sometime during the winter months of July to September. A map of its distributionis shown in Fig. 22.

Myrceugenia

105

37. Myrceugeniaoxysepala(Burret)Legrandet Kausel, in Legrand,Comun. bot. Mus. Hist. Nat. Montevideo 2(28): 5. 1953. Figs. 7E, 8J, 22. Luma oxysepala Burret,Repert. Spec. Nov. Regni Veg. 50: 51. 1941. ?Lumamacrosepala Burret,Repert. Spec. Nov. Regni Veg. 50: 53. 1941.Type. Herter26275, "Rio Grandedo Sul, Sao Franciscode Paula, Cimada Serra, 900 m" (holotype, B, apparently lost). ?Mvrceugeniamacrosepala (Burret)Legrandet Kausel, in Legrand,Comun. Bot. Mus. Hist. Nat. Montevideo2(28): 12. 1953. Myrceugenia macrocalyx [apparently should be macrosepala] var. obovata Mattos, Loefgrenia

65: 5. 1975. Type. Rambo s.n., Rio Grandedo Sul, "Vila Oliva p. Caxias," 12 Jul 1950 (holotype, PACA-47485).

Tree or shrub 2-6 m high; hairs whitish to reddish-brown,a mixtureof simple and dibrachiate;twigs greyish-whiteto reddish-brown,densely pubescent when young, glabrescentwith age; leaves glabrousto sparselypubescent (rarelydensely pubescent) below, glabrousto puberulentalong the midvein above, elliptic to obovate, 1.5-4 cm long, 0.6-1.5 cm wide, 2-2.6 times as long as wide; apex usually obtuse, sometimes acute or rounded;base cuneate to slightly acuminate; petiole channeled, sparsely to densely pubescent, 1-3 mm long, ca. 1 mm thick; midvein impressed for entire length or only proximallyabove, prominentbelow; lateral veins indistinctor up to ca. 10 pairs faintly visible; marginalveins equalling laterals in prominence; blades light to dark grey-green to reddish-brown above, pale yellow-greento lightbrownbelow, subcoriaceous,the marginslightly revolute, sometimes lighter than the rest of the blade; peduncles uniflorous,flattened, 1-3(-7) mm long, 0.5-1 mm wide, densely pubescent, solitary or in pairs in the leaf axils; bracteoles ovate to lanceolate, 4-7 mm long, 1.8-3.0 mm wide, ca. 2.5 times as long as wide, subcoriaceous, sparsely to densely pubescent without, less pubescent to glabrous within, loosely clasping the hypanthium;calyxlobes lanceolate to triangular,3.5-4.5 mm long, 1.5-3.0 mm wide, ca. 2 times as long as wide, subcoriaceous, densely pubescent without, sparsely pubescent to glabrous within, often slightly concave proximally; hypanthiumdensely pubescent, 1.4-2.0 mm long; disk 2-3 mm across, sparsely pubescent to glabrous, sometimes raised and conical; style 6-8 mm long, glabrous to puberulent;stamens 70-130, 4-8 mm long; anthers 0.3-0.6 mm long when dry; petals suborbicular to slightly elongate, 3-4 mm in diam.; ovary 3-4-locular; ovules 3-9 per locule; normalfruit unknown. Type. Loefgren s.n., "Brasilien: S. Paulo, Barreirodos Marins. Bluten weiss (Loefgren in herb. Mus. S. Paulo n. 3556 com. F. C. Hoehne)" (holotype, SP). Specimensexamined.BRAZIL. Parana:Mun.Tijucasdo Sul, Tabatinga,1 Dec 1964, Hatschbach 11920(MBM, MICH, MVM);Mun. Sao Jose does Pinhaes, Col. Roseira, 10 Oct 1966, Hatschbach 14814(NY). Santa Catarina:Mun. Bom Retiro, Riozinho, 1000m, 22 Jan 1957, Smith & Reitz 10280 (HBR, MVM, R, US); Bom. Jardim,CurralFalso, 1500m, 19 Mar 1959, Reitz & Klein8673; Mun. Sao Jos6 do Cerrito,Canoas, 1 Feb 1963,Reitz6492 (HBR, MVM,US); Mun. Lajes, ca. 18km south

of Lajes (ca. 27?50'S, 50?20'W), 1000 m, 22 Nov 1977, Landrum 2682, 2684 (MBM, MICH); Mun.

Urubici, MundoNovo, ca. 1500m, 12 Nov 1964,J. Mattos 12131(SP). Rio Grandedo Sul: Caxias, Vila Oliva, 15 Jul 1954, Rambo 55844 (MICH, MVM, PACA); Mun. Sao Franciscode Paula, 5 km along road to P6rto Alegre (ca. 29?30'S,50?15'W),ca. 1000 m, 26 Dec 1977, Landrum2947, 2951 (MICH, PACA); Mun. Cambara,Fortaleza, (ca. 29?S, 50?W),ca. 1100 m, 27 Dec 1977, Landrum 2981 (MICH, PACA).

Legrandhas used the name Myrceugeniaoxysepala for the type of that species and for what I believe to be three collections of M. acutiflorafrom Santa Catarina (1970). Otherwisehe has used the name M. macrosepala for what is here called M. oxysepala. It is possible that Burret's Luma macrosepala is a synonym of

L. oxysepala, but until type materialis found we cannot be sure. The possibility

Flora Neotropica

106 50

50 "'"

*..

0 ...: ::

FIG. 22. oxysepala.

--.;

< :'-

.. ....-

?'.

::*:'...

Distributions of Myrceugenia brevipedicellata, M. hoehnei, M. Ieptocalyx,

and M.

that the names are not synonyms is supported by the fact that Burret described both species in the same publication and in no place alludes to their similarity. Therefore, because there is still doubt about the true identity of L. macrosepala but no doubt about the identity of L. oxysepala, I have chosen to use the latter epithet. Myrceugenia oxysepala, more common in the planalto of southern Santa Catarina and Rio Grande do Sul than farther northward, is found in Araucaria forests and along the foggy eastern edge of the planalto. A map of its distribution is shown in Fig. 22. Myrceugenia oxysepala is most similar to M. leptocalyx and M. brevipedicellata. From the first it differs in having larger calyx-lobes and bracteoles. It is compared directly with the second in key K. Myrceugenia oxysepala flowers from December to March. 38. Myrceugenia leptocalyx Legrand, Comun. Bot. Mus. Hist. Nat. Montevideo Fig. 22. 2(28): 4. 1953 (name only); Darwiniana 11(2): 313. 1957. Mvrceugenia leptocalyx var. coriacea J. Mattos, Loefgrenia 65: 5. 1975. Type. Rambo s.n., Rio Grande do Sul, "Fazenda Englert p. Sio Francisco de Paula," 2 Jan 1955 (holotype, PACA56296).

Probably a shrub 1-3 m high; hairs a mixture of simple and dibrachiate, reddish-brown, the dibrachiate hairs usually quite asymmetrical; twigs densely pubescent when young, most hairs persisting until the first bark falls, sometimes arising in pairs in the leaf axils, the bark greyish or reddish-brown; leaves elliptic, 1-3.5 cm long, 0.6-1.5 cm wide, 1.4-2.7 times as long as wide, very sparsely pubescent to glabrous below, essentially glabrous above; apex acute to obtuse; base rounded to acute; petiole channeled, densely pubescent, 1-2 mm long, 0.5-

107

Myrceugenia

1 mm thick; midvein impressed for entire length or only proximally above, prominent below; lateral veins usually indistinguishable, rarely up to 5 pairs barely visible; marginal veins indistinct or faintly visible; blades light yellow-green to grey-green above, somewhat lighter below, coriaceous to submembranous, the upper surface lustrous or dull, the lower surface lustrous or dull, minutely wrinkled (at least when dry), sometimes appearing mottled; peduncles uniflorous, flattened, 1-3 mm long, 0.5-1 mm wide, densely pubescent, solitary or in pairs in the leaf axils; bracteoles ovate to lanceolate, 2-3 mm long, 1-1.5 mm wide, ca. 2 times as long as wide, subcoriaceous, sparsely to densely pubescent without, puberulent to glabrous within, clasping the hypanthium; calyx-lobes ovate to lanceolate, 2.5-3.5 mm long, 1.2-2.3 mm wide, ca. 1.5-2.5 times as long as wide, coriaceous, densely pubescent without, puberulent within, often slightly concave proximally within; hypanthium obconic, densely pubescent, 1.5-2 mm long; disk 2-2.5 mm across, pubescent to puberulent; ovary 3-4-locular; ovules 5-8 per locule; young flowers and mature fruit unknown. Type. A. Mattos & Labouriau s.n., Rio Grande do Sul, Sao Francisco de Paula, 13 Feb 1948 (holotype, MVM ex RB-63329). Myrceugenia leptocalvx is most similar to M. brevipedicellata and M. oxysepala, but can be distinguished from both by its bracteoles which are less than 4 mm long. Mvrceugenia leptocalyx is geographically separated from M. brevipedicellata and its leaves are mainly shorter and not as narrow as in that species. Calyx-lobe length serves to differentiate M. leptocalyx and M. oxysepala. In M. leptocalyx they are 2.5-3.5 mm long, whereas in M. oxysepala they are 3.5-4.5 mm long. Myrceugenia leptocalyx is known only from the two collections, from the area near Sao Francisco de Paula in Rio Grande do Sul. A map of its distribution is shown in Fig. 22. It probably flowers in January and judging from its apparent relatives M. leptocalyx is probably a shrub that grows along the borders between wet campos and Araucaria forests. Excluded Species Myrceugenia longipedicellata Barbosa Rodrigues, Myrt. Paraguay 2. 1903. Type. Hassler 5789, "Hab. in campis altoplanitie Yeruty, ad Serra do Maracayu, Paraguay" (holotype, G). This is definitely not a species of Myrceugenia. Dr. Rogers McVaugh believes that it is probably a species of Eugenia. Myrceugenia candolleana Legrand, Darwiniana 11(2): 314. 1957. New name based on Eugenia sellowiana A. P. de Candolle, proposed because Mvrceugenia sellowiana Berg (1857) already existed. This species may belong to Psidium. Two specimens (Glaziou 21178 at C and Duarte 2844 at MICH) which are similar to the photo of the holotype and which Legrand has referred to this species are at least superficially similar to P. incanum (Berg) Burret. Myrceugenia mosenii Kausel, Lilloa 33: 105. 1971. Type. Mosen 2840, Brazil, Sao Paulo, Santos, 15 Feb 1874 (holotype, S, n.v.; isotype, R). Although superficially similar to Myrceugenia reitzii, this is probably a species of Eugenia. It resembles closely specimens identified by Legrand as E. microcarpa Berg.

Flora Neotropica

108

Hybrids Interspecific hybridization is believed to be actively occurring between three different pairs of species of Myrceugenia. One of these cases has been studied previously (Landrum, 1974) and will be discussed only briefly here. Neither of the other two cases of hybridization has been specifically analyzed before, although hybridization has been mentioned by earlier workers as the possible explanation of some intermediate specimens. HI. Myrceugenia exsucca x ovata var. nannophylla Myrceugenia diemii Kausel, Revista Arg. Agron. 9: 239. 1942. Type. Diem s.n., "Rep. Argentina: Quetrihu6, no. 1146, Diem leg. (H. K.)" (lectotype, hereby designated, H-1091920; isolectotypes, H-1091921, H-1091922). Myrceugenia exsucca var. quetrihuensis Kausel, Lilloa 17: 53. 1949. Type. Diem 938, "Rep. Argentina: Lago Nahuel Huapi; Quetrihu6, Jos6 Diem leg. no. 938 (= no. 1192 Herb. Kaus.)" (lectotype, hereby designated, H-1091925; isolectotype, H-1091924). Specimens of M. exsucca x ovata var. nannophylla and highly introgressed specimens of M. exsucca examined. CHILE. Malleco: Parque Nacional de Tolhuaca, Laguna de Malleco, 4 Mar 1968, Zalenskv s.n. (H), 8-14 Mar 1968, Zalenskv s.n. (H). Osorno: Nadi entre Osorno y Puyehue, 19 Feb 1942, Kausel 1050 (H). ARGENTINA. Neuqu6n: Lago Nahuel Huapi, Quetrihu6, 20 Nov 1940, Diem 110 (LP, NY), 4 Feb 1940, Perez Moreau 35308 (LIL), 5 Mar 1942, Diem s.n. (H), 11 May 1942, Diem 362 (F, LIL), 12 May 1942, Diem 366 (H), 11 Feb 1947, Diem 998, 997 (H), 15 Mar 1947, Deim 999a 327, 330A (F), 2 Mar 1978, Landrum 3310, 3312, 3315, 3316, 3317, 3318, 3319, 3320, 3322, 3324 (LP, MICH); Lago Nahuel Huapi, Quetrihu6, Los Arrayanes, 19 Feb 1953, Boelcke & Correa 7033 (MICH), 1 Mar 1978, Landrum 3298, 3299, 3302, 3303 (LP, MICH); Lago Nahuel Huapi, Puerto Manzano, 1 Mar 1978, Landrum 3293, 3295, 3296, 3308 (LP, MICH). Rio Negro: Lago Nahuel Huapi, Puerto Pafuelo, 10 Apr 1945, Diem 941 (H), 3 Mar 1978, Landrum 3335 (LP, MICH). Specimens of typical Mvrceugenia exsucca used in this study but cited completely in the treatment of that species. Landrum 920, 3288, 3300, 3301, 3311, 3321, 3334, 3336; Diem 330B; Maldonado 21; Spegazzini 302-(No. 9); Kausel 619, 943, 4802, 5397, 5397a; Munoz 2761. Specimens of typical Myrceugenia ovata var. nannophylla used in this study but cited completely in the treatment of that taxon. Gunckel 3492; Junge 309; Landrum 929, 937, 3283, 3284, 3285, 3291; Philippi s.n.

The Kausel herbarium, now at H, contains the types of Myrceugenia diemii and M. exsucca var. quetrihuensis. There are duplicate specimens of each, so it has been necessary to designate lectotypes. Kausel (1942, 1949) implies that both Myrceugenia diemii and M. exsucca var. quetrihuensis could be hybrids. After examination of the types and several other collections from the Nahuel Huapi region, as well as possible hybrids from two localities in Chile, I concluded that hybridization was probably occurring. In order to study this apparent hybridization, extensive collections were made of both species and putative hybrids in the region of Lago Nahuel Huapi. This lake, located along the border between the provinces of Neuquen and Rio Negro in Argentina, very near to the border with Chile, is at about 780 m elevation. It extends westward well within the Andes and is surrounded by low hills and plains in the east. Rainfall at its western extreme is about 3 m per year while at its eastern extreme the yearly average is probably under one meter (Dimitri, 1972; Walter and Lieth, 1960-1967). One flower was taken from each collection made in this area and the stamens and ovules were counted; the length to width ratios of the bracteoles of the same flower were calculated and averaged; five leaves were measured and the average length calculated; the inflorescences were evaluated for the development of dichasia and the abundance of paired peduncles; and the density of the pubescence on the outer surface of the calyx-lobes was evaluated. In addition to the author's

109

Myrceugenia

.-?~~~~~~~~ !

. - .

. .ej'. "*~~~~~~~~~~~~~~~~~~~

.. * '-.6-'"

- CD as.;

G "-

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vey

enlty

~~~~~~~~~~~~~

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FIG. 23. Scatter diagram illustrating hybridization between Mvrceugenia exsucca and M. ovata var. nannophylla in the region of Lago Nahuel Huapi. See text for explanation of numerical values.

collections, three specimens from the same region but collected by other persons were also included in this study. None of them were types. The preliminary field observations indicated that hybridization was common in this area. There was a possibility that the parent species did not exist here in their pure form. In order to have an idea of what the unhybridized species would be like, five specimens of Myrceugenia ovata var. nannophylla and seven specimens of M. exsucca from areas in Chile where the species are not known to hybridize were also studied. The data for both of these groups of specimens were

110

Flora Neotropica

(

_..

;_s

-tb . E

.. Z

..::.... ..-'.-.:. ?

, .!-

j ..

::.~

: ,

FIG. 24. Map of Lago Nahuel Huapi Region in Argentina. The numbers indicate the following localities: 1-ca. 4 km north of El Cruce; 2-Villa Angostura; 3-east shore of Quetrihue Peninsula ca. 2 km south of base; 4-mallin (bog) on Quetrihue Peninsula; 5-Puerto Manzano; 6-Lago Patagua; 7-Los Arrayanes; 8-Puerto Anchorena; 9-Puerto Pafluelo; 10-Bahia L6pez. Wavy lines indicate lakes.

averaged and these values have formed the bases for idealized concepts of each taxon. All the information gathered in this study has been summarized in a scatter diagram (Fig. 23). Each small circle represents one specimen from the Nahuel ceugenia exsucca; the lower left-hand large circle, M. ovata var. nannophylla. The upper left-hand number within each circle corresponds to the collection locality on the map (Fig. 24). The upper right-hand number is the number of ovules in the ovary and in the lower half of the circle is the number of stamens. In a few cases data on ovules or stamens were not available so no number is shown. The arms on each circle represent the inflorescence and pubescence characters, while the axes of the diagram are the average leaf length and average bracteole ratio. By examination of the scatter diagram it is clear that there are many specimens

111

Myrceugenia

from Nahuel Huapi that are intermediate between typical Myrceugenia ovata var. nannophylla and M. exsucca and that these intermediates combine in several different ways the characteristics that normally separate these taxa. In other words there has been recombination of characteristics. This is exactly the pattern one would expect if the two species were hybridizing and the first generation hybrids were crossing among themselves or back-crossing with the parents. Collections used in the scatter diagram were made in ten different localities. If these are ordered geographically they stretch along a line that runs from NNW to SSE for ca. 36 km (Fig. 24). Those collections from the NNW extreme tend to be typical M. ovata var. nannophylla while at the SSE extreme the specimens seem to be typical M. exsucca or at least approach it closely. The change from west to east is the most important aspect of this line. Winds descending from the Andes become drier and warmer because of the adiabatic effect. Therefore conditions tend to be cooler and wetter towards the west and warmer and drier towards the east. What are the special conditions that allow hybridization to take place here? In other words, why are the hybrids more successful than either of the parents in some localities near Lago Nahuel Huapi? First let us examine how the parent species differ ecologically. Typical Myrceugenia ovata var. nannophylla grows from near Puerto Manzano (locality 5 in Fig. 24) and Villa Angostura (locality 2) to the north and west. Specimens that approach M. exsucca begin to appear about 2 km southeast of the base of the Quetrihue Peninsula (locality 3). The most typical specimens of M. exsucca studied in detail here were collected at Bahia Lopez (locality 10), Puerto Panuelo (locality 9) and Lago Patagua (locality 6). Thus M. ovata var. nannophyla seems to prefer the cool wet end of Lago Nahuel Huapi whereas M. exsucca seems to prefer a warmer, drier section of the lake. Myrceugenia exsucca is quite capable of growing with its roots submerged and seems to prefer such habitats. It apparently grows in its most typical form in sheltered conditions such as around small lakes and in bays on Lago Nahuel Huapi. It probably cannot grow in areas where waves are strong. Myrceugenia ovata var. nannophylla, at least in this region, grows farther away from the water's edge on the inland side of beaches. It also grows in more exposed areas than does M. exsucca. Perhaps its small leaves protect it from desiccating winds. The ecological characteristics of the two parent species and the intermediate condition of the hybrids are summarized below. M. ovata var. nannophvlla

M. exsucca

Grows beyond the water's edge on the inland side of beaches Has small leaves that may reduce desiccation by wind Grows at the cool wet end of lake

Grows with roots submerged or in saturated soil Has large leaves more vulnerable to desiccation by wind Grows in a warmer, drier area

Hybrid (M. diemiii) Grows in a more or less intermediate position, not at the water's edge and not as far inland as M. ovata var. nannophvlla Has leaves of an intermediate size Grows in intermediate conditions. Has been found with both parent species at the edges of their ranges.

The shores of Lago Nahuel Huapi, especially in relatively open areas, offer an intermediate habitat. For plants growing under these conditions, living at the water's edge may be impossible because of the destructive action of the waves

112

Flora Neotropica

in exposed areas. It is necessary for them to grow somewhat fartherinland than Myrceugenia exsucca normally does. The climate of much of the lake is drier and warmer than M. ovata var. nannophylla can apparentlytolerate. Then for the hybrid plants, the genes of M. ovata var. nannophyllaprovide the ability to live in exposed conditions, somewhatremovedfrom the lake shore; and the genes of M. exsucca provide tolerance to a relatively warm, dry climate. One curious fact is that Myrceugenia ovata var. nannophylla and M. exsucca

were never seen growing together at one locality. Myrceugeniaovata var. nannophylla was found with hybridsat Puerto Manzano(locality 5) and M. exsucca, or a somewhat introgressedform of it, was found with hybridsat Los Arrayanes (locality 7) and at locality 3. In the future both species may be found together, but perhaps their habitats do not quite meet and hybrids are the result of pollen being carried over short distances. In my own collections there were few specimens which conformedwell to the idealized concepts of Myrceugeniaexsucca based on the Chileanspecimens. This was not evident until the specimens had been thoroughlyanalyzed. Fortunately other specimens were available which confirmthe fact that M. exsucca exists in an apparentlypure state in this region. But if one looks at all the collections which have been made here, hybrids and apparentlyintrogressed specimens of M. exsucca outnumberapparentlypure specimens of M. exsucca. This may be because most collecting has been done around the large lake of Nahuel Huapi. I predict that the purest forms of M. exsucca will be found aroundthe small lakes on the QuetrihuePeninsulaand aroundthose near Puerto Pafuelo (locality 9). H2. Myrceugeniaexsucca x lanceolata Eugenia hridgesii Hooker et Arnott, Bot. Misc. 3: 322. 1833. Type. Cuming 99, "Chili" (holotype, perhaps K, n.v.; isotype, K). Eugenia multiflora [var.] / Hooker et Arnott, Bot. Misc. 3: 322. 1833. Specimen cited. Cuming 96, "Chili" (K). Eugenia pitra [var.] / angustifolia Hooker ex Berg, Linnaea 27: 265. 1856. Type. Cuming 96, "Chili" (holotype, perhaps W, n.v.; isotype, K). Myrceugenia bridgesii (Hooker et Arnott) Berg, Linnaea 30: 671. 1861.

Myrceugeniaexsucca var. bridgesii(Hooker et Arnott)Kausel, Lilloa 13: 142. 1948("1947").

Specimens of Myrceugenia exsucca x lanceolata examined. CHILE. Valparaiso: Valle de MargaMarga (ca. 33?10'S), Jaffuel & Pirion 3157 (GH); Valparaiso, Quebrada del Tranque, 18 Aug 1940,

Kausel 781 (F, H), Kausel 770, 777 (F); Algarrobo,Quebradadel Hotel Pacifico,4 Feb 1957,Kausel

4332; Algarrobo, Quebrada San Jer6nimo, 22-25 Mar 1951, Kausel 3216 (H). Colchagua: Antivero, 15 km de Aguas Buenas, 27 Nov 1938, Kausel 459 (H). Linares: Laguna Amargo, ca. 50 km east of Parral, ca. 720 m, 21 Feb 1972, all the following are members of a hybrid swarm which varies from pure or nearly pure M. exsucca to pure or nearly pure M. lanceolata, Landrum 910, 911, 912, 913, 914, 915, 916, 917, 918, 919, 922, 923, 926, 927, 928 (MICH).

Hybridization between Myrceugenia exsucca and M. lanceolata has been al-

luded to by Kausel (1948) and has been studied previously by Landrum(1974) throughanalysis of a hybrid swarm which was found aroundLagunaAmargoin the province of Linares in Chile. This locality, ca. 50 km east of Parralin the pre-cordilleraof the Andes, is unique. The small lake was formed by a landslide some time in the past. Water drains from the lake both above and below the ground. In the winter rainy season, enough water flows into the lake that water must flow out of the lake above the ground. In the summerdry season, the water level drops and all the water drainingout of the lake exits below ground. Thus the level of the lake fluctuates yearly by about 3-4 m. Myrceugeniaexsucca, as has been discussed before, is a tree of lake shores or swampy areas. It is quite capableof growingwith its roots submergedor in watersaturatedsoils.

Myrceugenia

113

Myrceugenialanceolata is a shrubthat mainlyinhabitsstreambanks. It appears to grow in areas with better drainagethan M. exsucca but probablycan withstand short periods of inundationas well as drought. Streams change with respect to the volume of water that flows in them, especially in an area of seasonal rains such as Chile. Aroundthe edge of LagunaAmargoexists a habitatthat is inundatedfor a few months of every year but also relatively dry for a few months. Hybrids between Myrceugenia exsucca and M. lanceolata are found all around the margin of the

lake, the variable habitat apparentlyfavoring them over either parent. Quite interesting, but perhaps not surprising,is the fact that two species of Sophora also hybridize aroundthe same lake (Donoso, 1974). Hybridization between Myrceugenia lanceolata and M. exsucca is not restrict-

ed to this area. Hybrids are rathercommon in the general vicinity of Valparaiso. Although I have not studied this area as extensively as that around Laguna Amargo, the following tentative explanation is offered. Both species grow in ravines and valleys of small streamsalong the coast. At the lower levels of these ravines and valleys M. exsucca is common, the habitat apparentlybeing wet throughoutthe year. At higher levels where inclines are steeper, M. lanceolata grows. Probablyhybridsgrow at an intermediateposition on the slopes or around pools which are full of water duringthe winter but fairly dry duringthe summer. This hypothesis is based on limited observations in the coastal zone of central Chile. Although Myrceugenia exsucca and M. lanceolata are undoubtedly best con-

sidered separate species, there are often specimens of both that appear to be slightly introgressed. Indeed the type of M. lanceolata would appearto be such a specimen and two of the author's collections (Landrum3446, 3451) cited under M. exsucca are probably not pure M. exsucca. They were collected in one of the coastal valleys described above, towards its upper, inland limit. H3. Myrceugenia euosma x glaucescens Myrceugeniaregnelliana var. dubia Legrand, in Legrandet Klein, Flora Ilustr. Catarinense [Mirt]:429. 1970. Type: Klein 3375, Santa Catarina,Lebon Regis, 900 m, 6 Dec 1962(holotype, MVM;isotype, HBR). Specimens of Myrceugenia euosma x glaucescens var. glaucescens examined. BRAZIL. Parana:

Mun. Sao Jose dos Pinhais,Col. Roseira, 30 Oct 1967,Hatschbach 17644(C, F, HB, MBM, MICH, MVM, RB, UC, US); Mun. Curitiba,Uberabade Baixo (ca. 25?30'S,49?20'W),ca. 1000m, 3 Nov 1977, Landrum2340, 2342 (MBM, MICH). Santa Catarina:Mun. Agua Doce, 28.5 km southeast of Horizonte, ca. 26?45'S,51?25'W,1000-1200m, 3 Dec 1964, Smith & Klein 13476 (H, HBR, R, SP, US); Mun. Bom Jardim,FazendadaLaranja,1400m, 13 Dec 1958, Reitz & Klein 4081 (H, MICH, NY, UC, US); Mun. Ca9ador,Fazenda Carneiros,northeastof Cagador,950-1100 m, 21 Dec 1956, Smith & Reitz 9011 (HBR, R, US); Mun. CampoAlegre, lower fazenda of Ernesto Scheide, ca. 900

m, 9 Nov 1956, Smith & Klein 7500 (HBR, R, US), 1 Feb 1957, Smith & Klein 10561 (HBR, MICH,

R, RB, US); Mun. Curitibanos,north of Curitibanos,850-950 m, 6 Dec 1956, Smith & Klein 8364 (HBR, US); Mun. Lajes, south of Lajes, km 24, ca. 900 m, 3 Dec 1956, Smith & Klein8166 (HBR, MICH, R, US); Matos Costa, P6rto Uniao, 1100m, 7 Jan 1962, Reitz & Klein 11705(US); Mun. Sao Joaquim, 1 km east of Bom Jardim,1100-1200m, 16 Jan 1957, Smith & Reitz 10197(HBR, US). Specimens of typical Myrceugeniaglaucescens used in this study but cited completely in the

treatment of that species. Rambo 45053, Rambo 30807, Schroeder s.n., Gallinal et al. 4504 1/3, Legrand 1708, Nicora 3229, Burkart 14301, Cabrera 3972.

Specimensof typical Myrceugeniaeuosma used in this study but cited completelyin the treatment

of that species. Smith & Klein 8468, 10557, 10564, 10606, 11043, 14971; Smith & Reitz 10178; Klein 3740; Hatschbach 17758.

Myrceugenia glaucescens var. glaucescens ranges from near Curitiba, Parana

to the vicinity of La Plata, Argentina.At the southernend of its range the plants are glabrousexcept for the hypanthiumand the leaves tend to be narrowwith an

Flora Neotropica

114

acute apex. Towards the north the plants tend to be sparsely pubescent on the young twigs, peduncles and calyx-lobes and the leaves are usually shorter and relativelybroadwith the apex bluntor rounded(Fig. 25 for leaf variation).Certain individualsin Santa Catarinaand Paranaare so pubescent and have the leaves so small that they are quite similarto other species, namely M. euosma and M.

ovata var. gracilis. Legrand and Klein (1970) suggest that these specimens are hybrids between M. glaucescens and M. ovata var. gracilis (=M. regnelliana) and

have namedthem M. regnelliana var. dubia. I agree with Legrandand Klein that they are hybrids, but I propose that they are the result of crosses between M.

glaucescens var. glaucescens and M. euosma. My interpretation is supported by

the fact that if one ranks the three species in question and the hybrids according to the average numberof ovules and average bracteole width, the hybridsfall

between M. glaucescens var. glaucescens and M. euosma, not between M. ovata var. gracilis and M. glaucescens var. glaucescens. The average values

for these characteristicsare listed below for each species and for the apparent hybrids taken as a group. Mean number of ovules per locule ? 0.97 . 7.73 + 0.74 8.67 + 1.34 9.57 + 0.92

M. ovata var. gracilis .....................................................4.08 M. glaucescens var. glaucescens ................................... Apparent hybrids ..................................................... M . euosm a ................................................................ Mean bracteole width in millimeters M. ovata var. gracilis .................................................... M. glaucescens var. glaucescens ........................................... Apparent hybrids ......................................... M. euosm a ................................................................

.........

0.48 1.02 . 1.24 1.62

+ 0.07 + 0.14 + 0.26 + 0.21

Anothercharacterthat indicatesthat Myrceugeniaeuosma is one of the parents of these hybrids is the pubescence on the lower surface of the leaves. Myrceugenia euosma has leaves that are densely pubescent beneath whereas in both M. glaucescens and M. ovata the leaves are glabrous or nearly so beneath. In the apparenthybrids the leaves are moderatelyto sparsely pubescent beneath. In Fig. 26, a scatter diagramis shown of nine specimens of Myrceugeniaeuosma, eight of M. glaucescens

var. glaucescens and six intermediates that are

believed to be hybrids. Each numericalcharacteris an average of a few separate measurementsor counts for each specimen. The pubescence charactersare an evaluation of each specimen. The intermediateposition of the suspected hybridsis illustratedin the diagram. It should be noted that they combine in various ways the characteristicsof the parents, which is consistent with the idea that they are F1hybridsor the products of subsequent crosses between the F1 hybrids or backcrosses with the parents. Collections of intermediatesand what appear to be highly introgressedspecimens of Myrceugeniaglaucescens are about as common as apparentlypure M. glaucescens from the planalto of Paranaand Santa Catarina.Thus hybridization is probablyfrequent there. Why are the hybrids successful in this region?I do not understandthe ecology of these species sufficiently to propose more than a tentative hypothesis. Myrceugenia glaucescens typically grows in lowland areas along rivers as far south as La Plata, Argentina.Perhapsit has migratednorthwardalong the Rio Uruguay in relatively recent geologic time. If it followed the tributariesof the Rio Uruguay into the planaltoof Parana,Santa Catarinaand Rio Grandedo Sul, it would have

115

Myrceugenia

0.,

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istics contributed by M, euosma might allow M,~~~~.' esen .o sriv . '~" sional drought, An extensive study of M, glauc4scens throughout its range be necessary before this hpothesis could be evalua,...

FIG. 25.

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Leaf variation in Mvrceugenia glaucescens var. glaucescens.

come in contact with M. euosma, a relatively xerophytic species common in the planalto. Moisture conditions along the small streams of the planalto are probably not as stable as along the larger rivers farther south. Thus xerophytic characteristics contributed by M. euosma might allow M. glaucescens to survive occasional drought. An extensive study of M. glaucescens throughout its range would be necessary before this hypothesis could be evaluated.

116

Flora Neotropica

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.::..:::... ~~~~P.' '"I ~~~~~~~~~~~~. '-:"? .-.::::"":. '~::.:.:.?.... ?-.?I?~ '~:::::'~:":"':~'""'"':' :" "::...""' :"'"::::::::::::::::::::: :-~:?' :::i:::: :!::: .;:::::-. '::.: ..,..":....'' -.'.'. '..x~: ... =.~...<:''::;" ":~"'... "' '"'.~ .... ......-.',-: . ?

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and dots with dark centers equal suspected hybrids. All numerical values are averages of a few measurements or counts.

117

Myrceugenia

[

FIG. 27. Selected specimens of Myrceugenia. A, M. rufescens (Hoehne s.n., 28 Sep 1931); B, M. campestris (Hatschbach 16471); C, M. exsucca (Hastings 474); D, M. lanceolata (Behn s.n., 4 Feb 1923).

118

Flora Neotropica

"' '-.

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28 eetdseieso FIG.~~~~~~~~~~~~~I

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FIG. 28. Selected specimens of Mvrceugenia. A, M. rufa (Landrum 3422); B, M. ovata var. ovata (Everdam 10656); C, M. obtusa (Merxmuller 24901); D, M. parvifolia (Hollermayer 587).

119

Myrceugenia h

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FIG. 29.

Illustration of Myrceugenia smithii and M. hatschbachii. A-C, M. smithii: A, young

branch, 1x; B, flower viewed from above afterthe petals and stamenshave fallen, 5x; C, flowerbud and peduncle, 5x (Smith & Klein 12411, type). D-F, M. hatschbachii: D, young branch, lx; E,

flower bud and peduncle, 5x; F, flower viewed from above after petals and stamens have fallen, 5x (Hatschbach 13043, type). Drawn by Mrs. Karen Douthit.

Flora Neotropica

120

? ,

...

'..

... ._

A .,

II? C

.-':-. . .. .......

FIG. 30. Selected specimens of Myrceugenia. A, M. leptospermoides (Merxmuller 24835); B, M. planipes (Hollermayer 1208); C, M. reitzii (Hatschbach 30532); D, M. kleinii (Hatschbach 16901).

121

Myrceugenia

~'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~P ".,. .~~: ~.~ .,*'~

~"''?i!~

.... ' ~'.~'~,.:~~~~~~~~~~~~~~~~~~~~~~~~~~ ~:.s,;~:.

?:.i:?;~~t:f?,

FIG.

31.

Selected

specimens

M~~~~~~~~~~~~~~~~~~~~~vrc4ugPnia. A, M. chrvsocarpa (Everdcrm

10551);

glau-~~~~~~~~

c4scens var. glauc4sc4ns(Schroeder s.n., Dec 1924); C, M. cucullura (Harschbuch 18633); M.~~~~~~~~~~, D,

(Du!{i! .";.!... s4rr'atorninosa~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

FIG. 31. Selected specimens of Mvrceugenia. A, M. chrvsocarpa (Everdam 10551); B, M. glaucescens var. glaucescens (Schroeder s.n., Dec 1924); C, M. cucullata (Hatschbach 18633); D, M. seriatoramosa (Dusen 13324).

Flora Neotropica

122

~~~~~A -~ qtiSM,0

z..

......

FI.32 elceseimn f ~rcueia M ocguni Lrde s. SO634 ,M Hrcbc 85,ioyeo .ln~cuclt,U) i,Mrnicni -J holotype

FIG. 32. Selected specimens of Myrceugenia. A, M. colchaguensis (Landbeck sn., SGO-64324, holotype); B, M. franciscensis (Hatschbach 18950, isotype of M. longipedunculata, UC); C, M. alpigena var. alpigena (Irwin 2778); D, M. alpigena var. rufa (Hatschbach 18158).

Myrceugenia

123

I4kI

FIG. 33. Selected specimens of Myrceugenia. A, M. acutiflora (Pessoal do Horto Florestal s.n., 22 Feb 1927); B, M. mvrcioides var. mnrcioides (Hatschbach 24377); C, M. mvrtoides (Pereira 6798); D, M. hrevipedicellata (Landrum 2830).

124

Flora Neotropica VII. ACKNOWLEDGMENTS

I am gratefulto all who gave me help and encouragementduringmy studies of Myrceugenia. My wife Sonia did many of the secretarialjobs related to this monograph and served as my field assistant in South America. Dr. Rogers McVaughalways kindly offered his time, and I have profitedon many occasions from his great experience with the Myrtaceaeand from his botanical knowledge in general. I thank the other members of my doctoral committee, Drs. William R. Anderson, WilliamS. Benninghoff,Peter B. Kaufman,and Arnold G. Kluge, for considering my dissertation and for their suggestions. I also appreciate the many helpful comments made by Dr. Rudolf Schmid of the University of California at Berkeley, who reviewed this monograph. I wish to thank those in charge of the herbariaof the following institutionsfor sending materialon loan for my study or for allowing me to consult their collections as a visitor: Royal Botanical Gardens, Kew; Museum National d'Histoire Naturelle, Paris; Conservatoireet JardinBotaniques, Geneve; Instituto Botanico dell'Universita, Torino; Botanische Staatssammlung, Miinchen; Naturhistorisches Museum, Wien; Botanical Museumand Herbarium,Copenhagen;Botanical Museum, University of Helsinki, Helsinki; Herbariumof the Departmentof Higher Plants, V. L. Komarov Botanical Institute, Leningrad;Gray Herbarium of Harvard University, Cambridge;The New York Botanical Garden, Bronx; Academy of Natural Sciences, Philadelphia;United States National Herbarium, Washington;Field Museumof NaturalHistory, Chicago;MissouriBotanicalGarden, St. Louis; University of California,Berkeley; University of Michigan,Ann Arbor; Museu Nacional, Rio de Janeiro; Jardim Botanico do Rio de Janeiro; HerbariumBradeanum,Rio de Janeiro;Herbario"Alberto Castellanos," Rio de Janeiro;Instituto de Botanica, Sao Paulo; Museu Botanico Municipal,Curitiba; Herbario"Barbosa Rodrigues," Itajai;HerbariumAnchieta, Sao Leopoldo; Museo Nacional de Historia Natural, Montevideo; Facultadde Agronomiay Veterinaria, Universidad Nacional del Nordeste, Corrientes; Fundacion e Instituto Miguel Lillo, Tucuman;Instituto de Botanica Darwinion, San Isidro; Museo de La Plata, La Plata; Museo Nacional de Historia Natural, Santiago; Facultad de Ciencias Forestales, Universidadde Chile, Santiago;Departamentode Botanica, Universidad de Concepcion, Concepcion; Instituto de Botanica, Universidad Austral, Valdivia. I am also grateful to the National Science Foundationfor providingfunds for field and herbariumstudies in South America and to the following persons and institutions for helping me with my field studies in South America: Dr. Pedro Carauta and Sr. Valerio Flechmann Ferreira of the Jardim Botanico, Rio de Janeiro;the Institutode Botanica, Sao Paulo; Dr. Gert Hatschbachof the Museu Botanico Municipal, Curitiba;Dr. Aloysio Sehnem of the HerbariumAnchieta, Sao Leopoldo; Dr. Elsa Zardiniof the Museo de La Plata, La Plata; the Museo Nacional de Historia Natural, Santiago;the Departamentode Botanica, Universidad de Concepcion, Concepcion;the Instituto de Botanica and the Facultadde Ciencias Forestales of the UniversidadAustral, Valdivia. Special thanks are due Dr. B. A. Krukoff. It has been his vision and financial backingthat have made possible my present position at the New York Botanical Garden, where this monographwent through the final stages of preparationfor publication. VIII. LITERATURE CITED Atchison, E. 1947. Chromosome numbers in the Myrtaceae. Amer. J. Bot. 34: 159-164.

Myrceugenia

125

Baker, P. E. 1967. An outline of the geology of the Juan Fernandez Archipelago. Geol. Mag. 104: 110-115. Berg, 0. 1855-1856. Revisio Myrtacearum Americae. Linnaea 27: 1-472. . 1857-1859. Myrtaceae. In: Martius, C. F. P. von, Flora Brasiliensis 14(1): 1-655. Bolkhovskikh, Z., V. Grif, T. Matvejeva & O. Zakharyeva. 1969. Chromosome Numbers of Flowering Plants. Academy of Sciences of the U.S.S.R., Leningrad [In Russian]. Briggs, B. G. & L. A. S. Johnson. 1979. Evolution in the Myrtaceae-Evidence from inflorescence structure. Proc. Linn. Soc. N.S.W. 102: 157-256. Burret, M. 1941a. Myrtaceen-Studien. Notizbl. Bot. Gart. Berlin-Dahlem 15: 479-550. . 1941b. Myrtaceenstudien II. Repert Spec. Nov. Regni Veg. 50: 50-60. Candolle, A. P. de. 1828. Myrtaceae. In: Prodromus Systematis naturalis regni vegetabilis 3: 207296. Carlquist, S. 1975. Ecological strategies of xylem evolution. University of California Press, Berkeley. Dimitri, M. J. 1972. La regi6n de los bosques andino-patag6nicos, sinopsis general. Instituto Nacional de Tecnologia Agropecuaria, Buenos Aires. Donoso, C. 1974. Un hibrido entre Sophora macrocarpa y Sophora microphylla. Fac. Cienc. Forest. Univ. Chile Bol. T6c. No. 30: 11-19. Estabrook, G. F. & C. A. Meacham. 1979. How to determine the compatibility of undirected character state trees. Math. Biosci. 46: 251-256. -, J. B. Strauch, Jr. & K. L. Fiala. 1977. An application of compatibility analysis to the Blackiths' data on orthopteroid insects. Syst. Zool. 26: 269-276. Fergulio, E. 1950. Descripci6n geol6gica de la Patagonia. Dir. Yac. Petrol. Fisc., Buenos Aires. Glaziou, A. F. M. 1905-1913. Plantae Brasiliae centralis a Glaziou lectae. Liste des plantes du Br6sil Central recueillies en 1861-1895. Mem. Soc. bot. France 1(3): 1-661. Gray, A. 1854. U.S. Expl. Exped., Phan.: 535-543. G. P. Putnam & Co., New York. Gunckel, H. 1972. Plantas chilenas descritas como nuevas por Juan Ignacio Molina y sus concordancias con la nomenclatura botanica actual. Mus. Nac. Hist. Nat. Noticiario Mensual (Santiago) 17(197): 3-11. Harrington, H. J. 1962. Paleogeographic development of South America. Bull. Amer. Assoc. Petr. Geol. 46: 1773-1814. Holmgren, P. K. & W. Keuken. 1974. Index Herbariorum. Osthoek, Scheltma and Holkema, Utrecht, Netherlands. Ingle, H. D. & H. E. Dadswell. 1953. The anatomy of the timbers of the south-west Pacific area. III. Myrtaceae. Aust. J. Bot. 1: 353-401. Johow, F. 1896. Estudios sobre la flora de las islas de Juan Fernandez. Cervantes, Santiago. Kausel, E. 1942. Contribuci6n al estudio de las Mirtaceas chilenas. Revista Argent. Agron. 9: 39-64. . 1944. Contribuci6n al estudio de las Mirtaceas chilenas, suplemento. Revista Argent. Agron. 11: 320-327. 1948 ("1947"). Notas Mirtol6gicas. Lilloa 13: 125-149. 1949. Notas Mirtol6gicas (suplemento). Lilloa 17: 51-55. .1956. Beitrag zur Systematik der Myrtaceen. Ark. Bot., ser. 2, 3: 491-516. Kluge, A. G. & J. S. Farris. 1969. Quantitative phyletics and the evolution of anuans. Syst. Zool. 18: 1-32. Landrum, L. R. 1974. Un hibrido entre Myrceugenia exsucca y Myrceugenia lanceolata. Fac. Cienc. Forest. Univ. Chile Bol. Tec. No. 30: 5-10. . 1980. A monograph of the genus Myrceugenia (Myrtaceae). Unpubl. Doctoral thesis, Univ. of Michigan, Ann Arbor. . 1981. The phylogeny and geography of Mvrceugenia (Myrtaceae). Brittonia 33: 105-129. Legrand, C. D. 1953. Nota preliminar sobre las especies de Mvrceugenia austrobrasilefas. Comun. Bot. Mus. Hist. Nat. Montevideo 2(28): 1-13. 1957. Representantes neotropicales del genero Myrceugenia. Darwiniana 11: 292-365. 1961. Mirtaceas del Estado de Santa Catarina (Brasil). Sellowia 13: 295-363. & R. M. Klein. 1970. Mvrceugenia. Flora Ilustr. Catar. [MIRT]: 333-453. Herbario "Barbosa Rodrigues," Itajai. McVaugh, R. 1968. The genera of American Myrtaceae-an interim report. Taxon 17: 354-418. Metcalfe, C. R. & L. Chalk. 1950. Anatomy of the Dicotyledons. Clarendon Press, Oxford. Meylan, B. A. & B. G. Butterfield. 1975. Occurrence of simple, multiple, and combination perforation plates in the vessels of New Zealand woods. New Zealand J. Bot. 13: 1-18. Navas, E. 1970. Distribucion geografica de las Mirtaceas Chilenas. Bol. Mus. Nac. Hist. Nat. Chile 29: 223-247. Niedenzu, F. 1893. Myrtaceae. In: K. Prantl and A. Engler, Nat. Pflanzenfam. 3(7): 57-105. Ragonese, A. M. 1976. Consideraciones sobre el problema de la clasificacion de los elementos tra-

Flora Neotropica

126

queales no perforadosde las dicotiled6neasy en especial de algunasMirtaceas.Darwiniana20: 476-490. Record,S. J. & R. W. Hess. 1943.Timbersof the New World.Yale Univ. Press, New Haven. Rutland, R. W. R., J. E. Guest & R. L. Grasty. 1965. Isotopic ages and Andean uplift. Nature 208:

677-678. Rye, B. L. 1979. Chromosomenumbervariationin the Myrtaceaeand its taxonomic implications. Aust. J. Bot. 27: 547-573. Sastrapradja,D. S. & C. Lamoureux. 1969. Variationsin wood anatomyof HawaiianMetrosideros (Myrtaceae).Ann. Bogor. 5: 1-83. Smith-White,S. 1948. Cytologicalstudies in the Myrtaceae.II. ChromosomeNumbersin the Leptospermoideaeand Myrtoideae.Proc. Linn. Soc. N.S.W. 73: 16-36. . 1950. Cytologicalstudies in the Myrtaceae.III. Cytology and phylogenyin the Chamaelaucoideae. Proc. Linn. Soc. N.S.W. 75: 99-121. . 1954. Cytologicalstudiesin the Myrtaceae.IV. The subtribeEuchamaelaucinae.Proc. Linn. Soc. N.S.W. 79: 21-28. Volkheimer,W. 1971. Aspectos paleoclimaticosdel TerciarioArgentino.Rev. Mus. Argent.Ci. Nat. "Bernardino Rivadavia," Paleontol. 1: 243-262. Walter, H. & H. Lieth. 1960-1967. Klimadiagramm-Weltatlas. Fischer, Jena.

IX. NUMERICAL LIST OF TAXA Myrceugenia 1. M. fernandeziana (Hooker et Arnott) Johow 2. M. rufescens (A. P. de Candolle) Legrand et Kausel 3. M. campestris (A. P. de Candolle) Legrand et Kausel 4. M. schultzei Johow 5. M. exsucca (A. P. de Candolle) Berg 6. M. lanceolata (Jussieu ex Jaume Saint-Hilaire) Kausel 7. M. rufa (Colla) Skottsberg ex Kausel 8. M. ovata (Hooker et Arnott) Berg a. var. ovata b. var. nannophylla (Burret) Landrum c. var. gracilis (Burret) Landrum d. var. acutata (Legrand) Landrum 9. M. pinifolia (F. Philippi) Kausel 10. M. obtusa (A. P. de Candolle) Berg 11. M. parvifolia (A. P. de Candolle) Kausel 12. M. leptospermoides (A. P. de Candolle) Kausel 13. M. hatschbachii Landrum 14. M. smithii Landrum 15. M. planipes (Hooker et Arnott) Berg 16. M. kleinii Legrand et Kausel 17. M. reitzii Legrand et Kausel 18. M. correifolia (Hooker et Arnott) Berg 19. M. chrysocarpa (Berg) Kausel 20. M. glaucescens (Cambessedes) Legrand et Kausel a. var. glaucescens b. var. latior (Burret) Landrum

21. M. scutellata Legrand 22. M. cucullata Legrand

23. M. seriatoramosa(Kiaerskou)Legrandet Kausel 24. M. colchaguensis (R. A. Philippi)Navas 25. M. franciscensis (Berg) Landrum

26. M. alpigena (A. P. de Candolle)Landrum a. var. alpigena

b. var. rufa (Berg) Landrum c. var. longifolia (Burret)Landrum 27. M. bracteosa (A. P. de Candolle)Legrand et Kausel 28. M. euosma (Berg) Legrand 29. M. acutiflora(Kiaerskou)Legrandet Kausel 30. M. venosa Legrand 31. M. myrcioides (Cambessedes) Berg a. var. myrcioides

b. var. acrophylla(Berg) Legrand 32. M. pilotantha (Kiaerskou)Landrum a. var. pilotantha

b. var. major (Legrand)Landrum 33. M. miersiana(Gardner)Legrandet Kausel

34. M. mvrtoides Berg

35. M. hoehnei (Burret)Legrandet Kausel

36. M. brevipedicellata (Burret) Legrand et

Kausel 37. M. oxvsepala (Burret)Legrandet Kausel

38. M. leptocalvx Legrand

127

Myrceugenia

X. LIST OF EXSICCATAE1 Almeida, J., 1265 (32a). Alt. Barb., 119 (23). Altamiro & Walter, 66 (8d). Anderson, W. R. et al., 36240 (26b). Anwandter, R., s.n. (7). Aravena, P., 18057 (15). Arechavaleta, J., s.n. (20a), Myrt. n 17 (34). Barba, R. de, 376 (5), 1003 (19), 1052 (19), 1756 (5), 1778 (5), 2283 (19). Barth, F190 (26a). Barreto & A. C. Brade, 16033 (8c). Barros, E., s.n. (5). Behn, s.n. (6), s.n. (11), 1175 (15). Behn, F., s.n. (15), 13658 (7). Behn, K., 23109 (10), 23110 (10). Bernasconi, s.n. (8b). Bernath, E., s.n. (5), s.n. (7), s.n. (8a), s.n. (9), s.n. (11), 871 (19), 886 (19), 1248 (7). Bertero, C. G., 1164 (6), 1167 (10), 1170(7), 1765 (7). Bertoni, 470 (28), 1999 (20a), 2398 (20a). Bettfreund & K6ster, 782 (20a). Bocher, T. W. et al., 1675 (5). Bock, C., 37 (1). Boelcke, 0., 2096 (5), 3913 (10), 3979 (18), 3983 (18), 3985 (18), 6465 (10), 6467 (18), 6503 (5), 6504 (5), 6517 (18), 6525 (7), 6529 (6), 6538 (6), 6539 (10), 6582 (18), 10548 (15), 10551 (8b). Boelcke, O. & Correa, 5834 (19), 7033 (HI), 7248 (8b). Bonpland, A., 1227 (20a). Brade, A. C., s.n. (2), 14608 (8c), 14609 (26a), 17264 (26a), 17295 (8c), 20053 (8c), 20333 (26a), 21290 (2). Braga, R. & H. Moreira, 507 (28). Bresolin, 801 (29). Bridges, s.n. (8a), 320 (5), 693 (15). Buchtien, O., s.n. (5), s.n. (11), s.n. (15). Burkhart, A., 3675 (20a), 4207 (20a), 4496 (20a), 4905 (20a), 8947 (20a), 14301 (20a). Burkhart, A. & J. C. Gamerro, 21840 (20a), 21844 (20a). Cabrera, A., 2510 (20a), 3972 (20a), 5031 (8b), 5038 (8b), 6006 (8b). Cabrera, A. R. Reitz & R. Klein, 9964 (8c), 9973 (17). Camargo, 0., 839 (8d), 1684 (8d), 2244 (8d). Campos Porto, P., 2423 (26a), 2424 (26a), 2716 (8c), 2787 (8c), 3423 (26a), 3424 (26a). Cantino, P., 40 (5), 80 (15). Capell, P., s.n. (36). Carrasco, 16 (12). Castellanos, A., s.n. (19), s.n. (20a), 24541 (21). Cecato, G. N., 16 (29). Chamisso, s.n. (7). Chapin, J. P., 1070 (4), 1071 (4). The three hybrids are designated by HI, H2, and H3.

Chebataroff, s.n. (20a). Claude Joseph, 953 (18), 963 (7), 2872 (10), 2913 (6), 3266 (11), 3304 (15), 3309 (11), 3319 (11), 3971 (8b), 4880 (8b), 5362 (15), 5396 (15), 5933 (12), 5936 (15), 5958 (10). Consigny, A., s.n. (19). Correa, 5610 (19). Costa, J. A. F., 9 (31a). Cuming, 30 (8a), 31 (8a), 96 (H2), 97 (6), 98 (6, 15), 99 (H2). Cumings, 102 (5). Cunningham, R. O., s.n. (15). Dawson, G., 928 (20a). Dawson & Schwabe, 2337 (5). Del Puerto, 3935 (34). Del Puerto & Marchesi, 3602 (34). Descole, H. R., 2599 (19). Dessauer, s.n. (7). Didrichsen, 3296 (6), 3339 (6), 3345 (10). Diem, J., s.n. (15), s.n. (H1), 93 (19), 106 (19), 110 (H1), 326 (5), 327 (H1), 330A (H1), 330B (5), 359 (8b), 362 (H1), 366 (H1), 938 (H1), 941 (H1), 997 (H1), 998 (HI), 999a (HI). Diogo, C., 414 (31b). Dionisio & Otario, 216 (29), 221 (31b). Dombey, J., s.n. (5), s.n. (10), s.n. (11), s.n. (12), 776 (9). Donoso, C. et al., s.n. (5), s.n. (10). Duarte, A. P., 4649 (32a). Duarte de Barros, 1069 (31b). Dusen, P., s.n. (12), 3318 (28), 3377 (8c), 3444 (31b), 8065 (33), 10263 (17), 13324 (23). Emygdio, 3085 (31b). Eskuche, U., 01470 (8d), 01471 (28), 1647b-4 (28). Eskuche, U. & Klein, 0100 (8b), 0282 (19). Eyerdam, W. J., 10519 (15), 10528a (15), 10551 (19), 10656 (8a). Ferrovia, s.n. (19). Fiebrig, 6472 (28). Friedrich, F., 1826 (5). Gallinal et al., 4504 (20a), 4504 1/3 (20a), PE 5188 (20a), PE 5580 (20a). Garaventa, A., 2145 (5), 2905 (5). Gardner, 421 (31a), 5712 (33). Gaudichaud, C., 234 (7), 1328 (34). Gay, C., s.n. (6), s.n. (11), s.n. (15). Gehrt, A., s.n. (32a), s.n. (35). Germain, s.n. (7), s.n. (10). Gillies, s.n. (6). Glaziou, A. F. M., s.n. (20b), 491 (32a), 1103 (32a), 2591 (32a), 5867 (31a), 6543 (32a), 6544 (26a), 6545 (8c), 8712 (32a), 13885 (31a), 13888 (29), 13891 (27), 13894 (29), 14271 (26b), 14821 (26b), 14833 (26b), 14834 (26a), 16050 (26a), 16052 (32a), 16066 (26b), 16073 (8c), 17003 (27), 17006 (23), 18249 (32a), 19363a (26b), 21147 (26c). Grandjot, C., 1503 (7), 3985 (12). Grisai, s.n. (18). Goes, O. C. & Octavio, 10 (32a). Grosse, 77 (15).

128 Gunckel, H., s.n. (15), 88 (15), 371 (5), 1453.4 (15), 1476.4 (15), 1941 (8a), 1959 (8a), 2131 (15), 3062 (15), 3492 (8b), 5054 (15), 14247 (5). Gurgel, s.n. (31a). Handro, O., 1056 (35). Harshberger, 885 (31a). Hastings, G. T., 248 (1), 474 (5). Hatschbach, G., 103 (33), 171 (28), 628 (31b), 909 (31b), 975 (33), 1132 (28), 1142 (8c), 1393 (23), 1744 (8c), 2226 (23), 2269 (31a), 2287 (28), 2433 (3), 2462 (31a), 3403 (8c), 3410 (28), 3412 (8c), 3418 (28), 3576 (28), 3579 (8d), 3581 (28), 4019 (33), 4021 (8c), 5050 (8c), 5634 (33), 6531 (32b), 6684 (26a), 6692 (8c), 6715 (22), 6785 (31b), 7311 (20b), 7336 (8c), 7355 (28), 7595 (8c), 7604 (31a), 7657 (31b), 7763 (31a), 8065 (31a), 8247 (2), 8780 (28), 9243 (17), 9258 (17), 9530 (31b), 9693 (8c), 9696 (26b), 9772 (31b), 9832 (31b), 9977 (33), 10035 (33), 10183 (3), 10337 (28), 10648 (28), 10968 (22), 10972 (31b), 10989 (31b), 10991 (23), 10995 (29), 11119 (31b), 11136 (31b), 11244 (33), 11342 (30), 11578 (16), 11646 (8c), 11920 (37), 12742 (3), 12797 (17), 12810 (31b), 13043 (13), 13482 (26b), 13483 (26b), 14546 (31b), 14593 (33), 14602 (31b), 14814 (37), 14997 (33), 15000 (31a), 15166 (20a), 15390 (28), 16116 (31b), 16242 (31a), 16423 (3), 16467 (33), 16471 (3), 16803 (31b), 16823 (8c), 16857 (8d), 16893 (3), 16896(31a), 16901 (16), 17182 (17), 17303 (3), 17562 (13), 17644 (H3), 17708 (8c), 17718 (28), 17758 (28), 17863 (31b), 18158 (26b), 18309 (2), 18446 (28), 18523 (31b), 18568 (8c), 18633 (22), 18678 (31a), 18950 (25), 19005 (31a), 19221 (31a), 20238 (17), 20598 (28), 20745 (8c), 22456 (17), 23240 (32b), 23405 (8c), 24377 (31a), 24660 (3), 24879 (17), 26205 (28), 26304 (28), 26403 (28), 27584 (8c), 29330 (31b), 30532 (17), 30670 (21), 31065 (26b), 30712 (21), 31810 (31b). Henz, E., s.n. (28), s.n. (34). Heringer, E. P., 9961 (26c). Herter, W. G., s.n. (20a), 91377 (20a). Hicken, C. M., s.n. (15). Hoehne, F. C., s.n. (2), s.n. (3), s.n. (32a). Hohenacker, R. F., 184 (11), 555 (8a). Hollermayer, P. A., s.n. (5), s.n. (11), 587 (11), 1207 (15), 1208 (15). Hunnewell, F. W., 16062 (19). Hutchinson, P. C., 193 (10). Imaguire, N., s.n. (31b), s.n. (33). Irwin, H. S., 2778 (26a). Irwin, H. S. et al., 28762 (27), 29383 (26b), 29386 (26b). Jaffuel, F., 1292 (10), 2929 (10). Jaffuel, F. & A. Pirion, 3157 (H2), 3170 (6). Jiles, C., s.n. (7), 5017 (7). Johow, F., s.n. (4), s.n. (5). Jonsson, G., 931a (28), 998a (2). Jozami, J. M., 256 (20a).

Flora Neotropica Junge, C., s.n. (19), 309 (8b), 852 (15), 3012 (12), 3015 (6). Kalela, A., 687 (8b), 713 (8b), 1165 (19), 1191 (19), 1228 (19), 1336 (5), 1449 (5), 1450 (8b). Kausel, E., s.n. (24), 273 (7), 347 (9), 446 (10), 447 (10), 458 (5), 459 (H2), 488 (6), 505 (10), 535 (19), 537 (19), 540 (19), 541 (8b), 542 (8b), 545 (19), 546 (19), 549 (8b), 551 (12), 557 (5), 561 (5), 562 (5), 567 (5), 579 (8b), 581 (8b), 589 (11), 596 (15), 599 (15), 602 (9, 12), 606 (5), 614 (11), 615 (11), 616 (12), 617 (11), 619 (5), 620 (5), 699 (6), 703 (5), 721 (6), 735 (5), 736 (5), 737 (5), 745 (11), 746 (9), 758 (10), 760 (5), 770 (H2), 777 (H2), 780 (6), 781 (H2), 782 (6), 783 (6), 784 (5), 846 (5), 850 (5), 906 (5), 907 (5), 908 (10), 920 (18), 925 (18), 926 (18), 933 (19), 934 (19), 943 (5), 996 (5), 997 (5), 1049 (8b), 1050 (HI), 1069(15), 1333 (8b), 1335 (8b), 1336 (8b), 1337 (8b), 1436 (11), 1590 (9), 1591 (9), 1592 (9), 1808 (10), 1834 (6), 1836 (6), 1840 (5), 1849 (10), 2370 (8b), 2373 (8b), 2419 (19), 2434 (19), 2441 (19), 2442 (8b), 2571 (7), 2584 (7), 2607 (7), 2612 (7), 2616 (7), 2639 (10), 2640(5, 12), 2702 (9), 2703 (6), 2728 (6), 2729 (6), 2730 (9), 2795 (15), 2796 (15), 2817 (8b), 2818 (8b), 3090 (7), 3216 (H2), 3514 (10), 3570 (9), 3628 (6), 3811 (18), 4055 (6), 4160 (18), 4182 (6), 4185 (7), 4327 (6), 4332 (H2), 4333 (5), 4341 (5), 4342 (5), 4350 (5), 4414 (7), 4552 (6), 4561 (18), 4624 (7), 4658 (7), 4673 (7), 4760 (15), 4799 (15), 4802 (5), 4882 (8b), 4911 (18), 4978 (8b), 5019 (7), 5381a (7), 5384 (6), 5385 (6), 5397 (5), 5397a (5), 5461 (5). Kiehl & Franco, s.n. (20b). King, s.n. (8a). Klein, R., 72 (17), 112 (3), 490 (31a), 528 (17), 567 (31b), 828 (17), 1187 (31b), 1196 (31b), 1366 (31a), 1482 (31b), 1518 (31b), 1540 (16), 1546(17), 1579(16), 1600(3), 1606(17), 1610 (17), 1925 (30), 1934 (31b), 1988 (31a), 2905 (31a), 3375 (H3), 3388 (8c), 3448 (8c), 3625 (8c), 3635 (8c), 3740 (28), 3761 (28), 3968 (8c), 4010 (8c), 4807 (31b), 7830 (32a), 7936 (8d), 8255 (29), 9648 (29). Klein, R. & Bresolin, 7717 (32a), 7771 (32a), 7916 (8d), 8173 (29), 9765 (8d), 9846 (8d), 10624 (21). Klein, R. & Souza, 7324 (29), 8079 (31b). Krapovickas, A. et al., 16802 (20a). Krause, s.n. (5), s.n. (8a). Kroll, H., s.n. (5). Kuhlmann, J. G., s.n. (26a), s.n. (32a). Kuhlmann, M., s.n. (31b), s.n. (33), 2185 (8c). Kuhlmann, M. & E. Kuehn, s.n. (31b). Kummrow, 355 (28). Kunkel, G., s.n. (15), M328 (5), 407 (10). Landbeck, s.n. (24). Landrum, L. R., 871 (8b), 876 (8a), 888 (11), 900 (8a), 910-923 (hybrid swarm of M. exsucca and M. lanceolata), 925 (6), 926 (H2), 928 (H2), 929 (8b), 935 (8b), 937 (8b), 938 (10),

Myrceugenia 939 (8a), 940 (8a), 944 (7), 945 (7), 946 (5), 947 (15), 2004 (31a), 2021 (32a), 2023 (32a), 2067 (31a), 2073 (31a), 2086 (33), 2110 (26a), 2111 (27), 2112 (26a), 2192 (29), 2205 (29), 2216 (32a), 2235 (33), 2274 (3), 2275 (3), 2305 (31a), 2312 (31a), 2318 (31a), 2340 (H3), 2341 (20a), 2342 (H3), 2370 (32b), 2393 (29), 2404 (33), 2407 (29), 2408 (33), 2409 (30), 2410 (30), 2411 (30), 2431 (28), 2433 (28), 2434 (28), 2455 (8c), 2458 (28), 2465 (20a), 2466 (20a), 2545 (13), 2552 (13), 2553 (13), 2554 (13), 2556 (13), 2617 (14), 2618 (14), 2619 (14), 2620 (14), 2621 (14), 2622 (14), 2623 (14), 2624 (14), 2625 (14), 2633 (32b), 2637 (8c), 2648 (28), 2649 (32b), 2650 (32b), 2651 (32b), 2652 (32b), 2653 (28), 2658b (20a), 2659 (20a), 2666 (22), 2671 (22), 2680 (28), 2681 (20a), 2682 (37), 2684 (37), 2712 (28), 2717 (28), 2718 (28), 2719 (28), 2722 (26b), 2723 (26b), 2730 (26b), 2798 (27), 2815 (8c), 2817 (27), 2819 (27), 2820 (36), 2821 (36), 2823 (33), 2824 (27), 2825 (27), 2826 (8c), 2827 (8c), 2828 (8c), 2830 (36), 2831 (27), 2849 (32a), 2857 (31b), 2858 (31b), 2859 (31b), 2860 (31b), 2861 (31b), 2862 (31b), 2866 (31b), 2872 (17), 2873 (3), 2874 (3), 2874 (3), 2875 (31b), 2876 (17), 2880 (3), 2883 (31b), 2890 (31a), 2906 (29), 2910 (32b), 2914 (31a), 2921 (30), 2923 (31a), 2931 (29), 2932 (30), 2933 (30), 2934 (31a), 2942 (33), 2946 (33), 2947 (37), 2949 (33), 2951 (37), 2953 (22), 2960 (22), 2961 (26b), 2962 (26b), 2965 (28), 2972 (32b), 2976 (32b), 2978 (33), 2981 (37), 2982 (22), 2984 (8c), 2986 (28), 3001 (31b), 3004 (22), 3005 (28), 3012 (8d), 3013 (8d), 3014 (8d), 3023 (34), 3024 (34), 3025 (34), 3043 (34), 3044 (34), 3045 (34), 3046 (34), 3084 (5), 3086 (9), 3090 (9), 3092 (9), 3093 (9), 3106 (15), 3107 (10), 3110 (5), 3112 (6), 3113 (10), 3123 (15), 3124 (5), 3125 (11), 3127 (11), 3137 (5), 3143 (15), 3163 (5), 3164 (11), 3165 (15), 3169 (8b), 3172 (8b), 3175 (8b), 3176 (8b), 3177 (15), 3178 (15), 3179 (15), 3181 (8a), 3184 (8a), 3188 (11), 3192 (11), 3192 (11), 3194 (8a), 3195 (8a), 3205 (8a), 3206 (8a), 3213 (8a), 3232 (5), 3242 (5), 3271 (11), 3273 (5), 3283 (8b), 3284 (8b), 3285 (8b), 3288 (5), 3290 (8b), 3291 (8b), 3292 (8b), 3293 (H1), 3294 (8b), 3295 (H1), 3296 (H1), 3298 (H1), 3299 (H1), 3300 (5), 3301 (5), 3302 (H1), 3303 (H1), 3308 (HI), 3310 (H1), 3311 (5), 3312 (H1), 3315 (H1), 3316 (5), 3317 (H1), 3318 (H1), 3319 (H1), 3320 (H1), 3321 (5), 3322 (H1), 3324 (H1), 3334 (5), 3335 (H1), 3336 (5), 3341 (19), 3342 (19), 3343 (19), 3347 (19), 3363 (10), 3379 (9), 3381 (9), 3384 (9), 3386 (9), 3421 (10), 3423 (18), 3424 (7), 3425 (7), 3427 (7), 3436 (18), 3439 (18), 3442 (5), 3446 (5), 3451 (5), 3454 (24), 3455 (24), 3456 (24), 3457 (24), 3458 (24), 3459 (24), 3460 (24). Landrum, L. R. & C. Donoso, s.n. (5), s.n. (6), s.n. (9), s.n. (10), s.n. (11).

129 Lanstyak, L., 252 (26a). Laroche, R. & J. Almeida, 1365 (32a). Lechler, W., 563 (5), 868 (11), 875 (15). Lee, L., s.n. (15). Legrand, C. D., 139 (20a), 684 (28), 724 (20a), 734 (28), 754 (28), 815 (20a), 1387 (20a), 1613 (20a), 1616 (20a), 1708 (20a), 2117 (28), 2210 (20a), 2211 (20a), 2485 (20a), 2486 (20a), 2488 (20a), 2492 (28), 3480 (28), 4183 (20a), 4184 (20a). Lindemann, J. C. & J. H. de Hass, 1892 (33). Loefgren, A., s.n. (8c), s.n. (31a), s.n. (37). Looser, G., 2574 (5), 2732 (19), 2733 (8b), 4360 (5). Lourteig, A., 2266 (28). Luederwaldt, H., s.n. (2), s.n. (8c). Lund, II-39 (27). Lutzelburg, 6550 (8c). Lycinger, E., 999 (5). Magalhaes, s.n. (27). Mahu, M., 2014 (8a). Maige, W., s.n. (6). Maldonado, R., 21 (5). Marticorena, C. et al., 894A (5), 1053 (12), 1070 (12). Marticorena, C. & Furet, 83 (19). Marticorena, C. & Matthei, 495 (24). Martinez Crovetto, R., 3225 (5). Martinez Crovetto, R. & A. Leguizamou, 5405 (20a). Martins, H. F., 290 (3 la). Martius, K. F. P. von, s.n. (2), s.n. (3), s.n. (20a), s.n. (25), s.n. (26a), s.n. (27). Mattos, A. & L. Labauriau, s.n. (20a), s.n. (28), s.n. (31b), s.n. (33), s.n. (38), 288 (22). Mattos, J., s.n. (32a), 3916 (28), 8869 (31a), 8945 (33), 9069 (3), 9102 (3), 10568 (32a), 10569 (32a), 11461 (35), 11719 (35), 11770 (3), 11807 (31a), 11915 (28), 12131 (37), 13491 (31a), 13993 (25), 14027 (2), 14376 (8c), 14736 (36), 14838 (35), 14888 (2), 14900 (8c), 15051 (8c), 15385 (8c), 15386 (27), 15389 (31a), 15487 (20b), 15623 (35), 15776 (33), 15798 (8c), 15800 (27), 15905 (27). Mertens, s.n. (7). Merxmiiller, H., 24835 (12), 24901 (10). Mexia, Y., 4257 (33), 8023 (15). Meyer, F., 9442 (4), 9509 (1), 9527 (1), 9709 (10). Meyer T., s.n. (19), 7363 (19), 9304 (5). Milano, s.n. (5). Montero, G., 1288 (6), 1335 (15), 1337 (8a), 2206 (19), 3192 (7), 3511 (10). Morrison, J. L., 16731 (10), 17601 (5). Morrison, J. L. & R. Wagenknecht, 17449 (19). Moseley, H. N., s.n. (1). Movia, C., s.n. (8b). Mufioz, C., 2756 (5), 2761 (5). Mufioz, C. & Johnson, 2724 (10). Neger, F. W., s.n. (15). Nicora, E. G., 3229 (20a), 3950 (5), 6384 (20a), 6386 (20a). Occhioni, 256 (31a). Osorio, H., 32 (20a).

130 Pabst, E. G., 6205 (8d). Palacios, M. A., 13 (20a). Pavon, J. A., s.n. (11), 890 (10). Pereira, E., 20 B (26a), 6243 (3 lb), 6798 (34), 7691 (28), 8565 (28). Pereira, E. & A. P. Duarte, 4522 (29). Perez Moreau, R. A., s.n. (5), s.n. (8b), 573 (5), 35308 (H1). Pedersen, T. M., 1544 (8b), 1607 (19). Pessoal do Horto Florestal-Rio de Janeiro, s.n. (26b), s.n. (29), s.n. (31a). Pfister, A., 4810 (12), 4821 (8a), 18543 (24). Philippi, R. A., s.n. (5), s.n. (8a), s.n. (8b), s.n. (9), s.n. (11), s.n. (12), s.n. (15), s.n. (19), 69 (8a), 136 (5), 667 (18), 727 (5), 728 (10), 732 (10). Pickel, B., 5374 (2). Pisano, E. & R. Baraona, 1393 (10). Pisano, E. & D. Geni, 1273 (11). Pisano, E. & A. Montaldo, 1421 (1), 1444 (1), 1470 (1). Plaumann, F., 275 (22). Poeppig, E. F., 76 (10), 114 (7), 143 (7), 144 (6), 421 (12), 456 (15), 457 (15), 926 (19). Pohl, 1013 (2). Poulsen, E. M., s.n. (5), s.n. (6), s.n. (15), 190 (6). Quezada, M. et al., 2 (9). Rambo, B. (numbers apparently equal those of PACA), 4319 (22), 26094 (34), 30795 (28), 30805 (20a), 30807 (20a), 38259 (34), 38694 (34), 39039 (34), 39588 (34), 39714 (34), 40298 (20a), 41820 (34), 42654 (3), 45053 (20a), 45444 (8d), 45706 (20a), 46640 (34), 47189 (28), 47485 (37), 49372 (28), 49649 (28), 50046 (28), 51648 (28), 51669 (20a), 53920 (32b), 53979 (22), 54062 (26b), 54094 (26b), 54483 (8d), 55844 (37), 55958 (8d), 56280 (26b), 56296 (38), 56364 (31b), 56578 (20a), 56597 (20a). Reed, E. C., s.n. (15). Regnell, A. F., I no. 130 (31b), II no. 120 (8c), III no. 570 (20b). Reiche, C., s.n. (18), s.n. (24). Reineck, E. M. & J. Czermak, 163 (34), 263 (34). Reitz, R., 1978 (28), 1985 (8c), 2413 (8c), 2420 (28), 2425 (8c), 2426 (28), 2574 (8c), 2576 (8c), 2741 (8c), 2744 (31b), 2769 (8c), 2770 (28), 3577 (28), 3651 (28), 3697 (28), 4398 (31b), 5192 (28), 5474 (30), 5476 (33), 5483 (31b), 6492 (37). Reitz, R. & R. Klein, 1292 (3), 1551 (31b), 1837 (31a), 1901 (31a), 2028 (17), 2122 (17), 2400 (31b), 2581 (31b), 2875 (31a), 3595 (31b), 4081 (H3), 4295 (31a), 4451 (31b), 4769 (32b), 4780 (2), 4788 (2), 4912 (28), 4986 (3), 5191 (2), 5192 (8c), 5197 (8c), 5431 (32b), 5942 (22), 5991 (30), 6024 (26b), 6030 (32b), 6051 (8c), 6110 (32b), 6160 (26b), 6280 (31b), 6435 (26b), 6586 (31b), 6645 (33), 7199 (8c), 7224 (8c), 7368 (14), 7427 (8d), 7741 (28), 7774 (8c), 7807 (8d), 7834 (28), 7870 (8c), 8093 (32b), 8131 (28), 8393 (26b), 8403 (32b), 8673

Flora Neotropica (37), 8675 (26b), 8760 (33), 9007 (31a), 9544 (32b), 9672 (3 la), 9677 (3 la), 9724 (3 la), 9733 (32a), 10417 (8c), 10466 (8c), 10490 (8c), 10496 (26a), 10592 (8c), 10673 (31b), 10756 (3 la), 10775 (32b), 10835 (3 la), 10846 (3 la), 10992 (32a), 10993 (8c), 11705 (H3), 11750 (31a), 11937 (21), 12430 (31b), 13528 (20b), 13535 (20b), 13735 (8c), 16491 (31b), 17790 (28). Reuthell, I. V., s.n. (5). Ricardi, M., 7474 (6), 10112 (24). Ricardi, M., C. Marticorena & Matthei, 1214 (19). Riedel, L., s.n. (26b), s.n. (31b), s.n. (33). Robello, s.n. (2). Rodrigo, A. P., 2219 (20a). Rodrigues, 578 (20a), 644 (10), 695 (20a). Rojas, T., 5995 (28), 6054 (28), 7901 (28), 9609 (28). Roivainen, H., 3218 (5). Rosengurtt et al., A 799 (20a), B 799 (20a), B 2582 (28), B 2743 (20a), B 4011 (20a), PE 4532 (20a), B 4801 (28), B 4801 1/2 (28). Rudolph, C., 4578 (19). Ruiz, H. & J. Pavon, s.n. (11). Rusby, H. H., 582 (6), 584 (6). Saelzer, F., s.n. (5). Saint-Hilaire, A., s.n. (27), s.n. (31a). Saldanha, 5233 (31b). Sargent, C. S., s.n. (5), s.n. (15), s.n. (19). Scala, A. C., 271 (20a), 272 (20a), 273 (20a), 274 (20a), 275 (20a), 276 (20a). Schajouskoy, S., s.n. (5). Schazmann, s.n. (8a). Schinini, A., 4358 (28). Schlegel, F. & A. Consigny, s.n. (18). Schroeder, s.n. (20a), s.n. (28), s.n. (34). Schwabe, G. H., 10 (15), lOa (15). Schwarz, 3958 (28). Seki, T., 350 (15). Sellow, F., s.n. (2), s.n. (3), s.n. (20a), s.n. (26b), s.n. (27), s.n. (28), s.n. (34). Silveira, H., s.n. (31a). Silverira, A., s.n. (8c). Simon, J. P., 233 (18). Skottsberg, C., s.n. (4), 75 (1), 134 (4), 816 (18), 1478 (10). Smith, L. B., 1734 (8c). Smith, L. B. & R. Klein, 7361 (8c), 7385 (8c), 7500 (H3), 7507 (31a), 7524 (28), 7793 (8c), 7835 (8c), 7842 (8c), 7863 (32b), 7922 (8c), 8022 (32b), 8113 (28), 8166 (H3), 8231 (28), 8232 (20a), 8364 (H3), 8369 (28), 8393 (31b), 8394 (8c), 8468 (28), 8494 (35), 8499 (31a), 8519 (8c), 8533 (8c), 9649 (28), 10557 (28), 10561 (H3), 10563 (26b), 10564 (28), 10606 (28), 10614(28), 11043(28), 11326(22), 11366 (20a), 11469 (28), 11525 (20a), 11540 (28), 11673 (31b), 12006 (17), 13072 (20a), 13122 (20a), 13351 (20a), 13366 (31a), 13368 (20a), 13370 (20a), 13476 (H3), 13805 (20a), 14035 (31b), 14156 (28), 14344 (26b), 14971 (28). Smith, L. B. & R. Reitz, 8573 (28), 8834 (31b),

131

Myrceugenia 8921 (26b), 8955 (8c), 8970 (28), 9011 (H3), 9014 (8c), 9822 (20a), 9873 (31b), 9877 (28), 9943 (28), 10178 (28), 10195 (28), 10197 (H3), 10280 (37), 10466 (32b), 12411 (14), 12844 (28), 14281 (26b). Smith, L. B., R. Reitz & L. Caldato, 9570 (28). Smith, L. B., R. Reitz & R. Klein, 7688 (28), 7729 (32b), 7731 (8c), 7732 (8c), 7932 (31b), 7733 (8c), 10297 (8c). Smith, L. B., R. Reitz & O. Sufridini, 9484 (28), 9486 (20a). Sobrinho, 505 (27). Solbrig, 0., 3711 (4), 3730 (4), 3809 (1). Sparre, B., 2308 (10), 3953 (8a). Sparre, B. & L. Constance, 10677 (19), 10839 (11), 10861 (5). Sparre, B. & Planella, 140 (1), 198 (1). Spegazzini, R. A., s.n. (8b), 302 no. 9 (5). Strang, 225 (26a). Sucre, D., 2325 (8c), 2462 (32a), 2539 (33), 4655 (26a), 4677 (8c), 7567 (32a). Sucre, D. et al., 7273 (2).

Tessmann,G., 6115 (33). Thaxter, s.n. (15). Ule, E., 706 (31b), 918 (31b), 2493 (26b). United States ExploringExpedition-C. Wilkes, s.n. (6), s.n. (7), s.n. (10), s.n. (18). Usteri, P. A., s.n. (2). Van Emelen, A., 19 (20b). Veloso, H., 16b(31b). Vidal, 97 (26a), II-705(26a). Villagran& Tapia, s.n. (18). Wagenknecht,R., s.n. (18). Warming,J. E. B., s.n. (27). Wawra,II 483 (26a). Weber, s.n. (1), s.n. (4). Werdermann,E., 299 (11), 900 (18), 1248 (19), 1258 (19). West, J., 4831 (19), 4861 (15), 4884 (8a), 5079(10), 5192 (10).

Widgren,J. F., 556 (8c). Zalensky, s.n. (HI). Zollner,0., 7129 (7), 7162 (6).

132

Flora Neotropica XI. INDEX OF SCIENTIFICNAMES

Note: Page numbers in boldfaceindicate primarypage references, page numbers with * indicate pages with illustrationsor maps. Actinodium 18 Amomyrtus luma 18, 31 Araucaria 11*, 46, 54, 73, 79, 80, 86, 99, 100, 104, 107 Baeckea 18 Beaufortia 18 Blepharocalyx 5, 6, 7, 18 gigantea 18 tweediei 18 Britoa 5 Callistemon 18 Calycolpus 5 Calycorectes 5 Calyptranthes 2, 5, 6 Campomanesia 5, 18 Darwinia 18 Eucalyptus 18 citriodora 18 globulus 18 Eugenia 2, 5, 17, 18, 19, 60, 107 acrophylla 94, 95 acutiflora 86 alpigena 76, 78 anceps 81 aprica 84, 86 araujoana 66, 69 bagensis 66 bagensis var. angustifolia 66 bagensis var. avenia 67 bagensis var. latifolia 67 brachymischa 81, 82, 83 brachymischa forma pedunculata 81, 83 bracteosa 81, bridgesii 112 buxifolia 62, 63 cambessedeana 66 campestris 33 canelonensis 67 chrysocarpa 62 colchaguensis 74 corralensis 35 correaefolia 61 cumingii 43 distans 33 distoma 56, 57 dombeyana 38 elegans 67 estrellensis 72, 94 euosma var. euosma 84 euosma var. lutescens 83, 84 euosma var. rufescens 84

expallens 81 exsucca 35 exsucca var. apiculata 35, 37 exsucca var. exsucca 35 exsucca var. patagua 35 exsucca var. peruviana 35 exsucca var. temu 35 fernandeziana 30 ferruginea 40 franciscensis 75 fuliginea 79 fuliginea var. rufa 79 glaucescens 63, 66 gudilla 38 hypericifolia 92 ibitipocensis 81 itatiaiensis 45 lanceolata 38 leptospermoides 51 leptospermoides var. latifolia 51 leptospermoides var. leptospermoides 51 leptospermoides var. longifolia 51 leptospermoides var. microphylla 51 lumilla 30 maritima 61 microcarpa 107 miersiana 99 miersiana var. costata 99 miersiana var. glomerata 99 miersiana var. membranacea 99 miersiana var. miersiana 99 miersiana var. venosa 99 montana 81, 82, 83 multiflora 35 multiflora var. / 112 myrcioides 91, 92 nana 84 nana var. congesta 84, 86 nana var. effusa 84, 86 nana var. nana 84 obtusa 49 obtusiflora 92 ovata 42, 43 pallida 67, 69 parvifolia 50 patagonica 62 petiolata 62 philippii 62 pilotantha 95, 97 pinifolia 47 pitra 35 pitra var. angustifolia 112 planipes 56, 57 planipes var. genuina 56 planipes var. grandifolia 56 planipes var. planipes 56 planiramea 32

133

Myrceugenia poeppigiana 32 polyantha 49 raran 49 regnelliana 45 ribeireana 67 rufa 40 rufescens 32 sellowiana 107 seriato-pedunculata 79 seriato-ramosa 73 stenophylla 38 stenophylla var. angustifolia 38, 39 stenophylla var. latifolia 38, 39 sticheromischa 79 temu 35, 37 thalassaia 61 thymifolia 51 trichocarpa 43 Eugeniinae 4*, 5, 6, 7 Eugenioideae 5 Gomidesia 2, 5, 6 Hexachlamys 5 Hypocreales 16 Homoranthus 18 Leptospermaceae 4 Leptospermoideae 4 Leptospermum 18 Luma 2, 4*, 6, 7, 18, 19, 69, 86 acrophylla 94 acutiflora 86 alpigena 78 angustior 67 apiculata 17, 18, 19, 57 aprica 84 araujoana 67 baeckeoides 50 bagensis 67 bracteosa 81 brevipedicellata 104 cambessedeana 82 campestris 33 canelonensis 67 chrysocarpa 62 cinerea 84 cinnamomeotomentosa 101 corralensis 36 correaefolia 61 cumingii 43 distans 33 dombeyana 38 elegans 67 euosma 84 estrellensis 94 expallens 81 exsucca 35 fernandeziana 30 ferruginea 40 filibracteata 103 fuscovelutina 99 glaucescens 67

gracilis 45, 46 hoehnei 103 itatiaiensis 45 latior 69 longifolia 81 macahensis 97 macromischa 45 macrosepala 105, 106 mesomischa 73 miersiana 99, 103 mucronata 75 myrcioides 92, 103 myrtoides 101 nana 84 nannophylla 44 obtusa 48 obtusiflora 92 oreophila 45 ovata 43 oxysepala 105, 106 pallida 67 philippi 62 pilotantha 97 pitra 35 planiramea 32 poeppigiana 32 regnelliana 45 rufa 40 rufescens 32 schultzei 34 seriato-pedunculata 79 seriato-ramosa 73 stenophylla 38 sticheromischa 79 temu 35 ulei 94, 95 Marlierea 2, 5, 6 Metrosideros 18 Myrceugenella 2 Myrceugenia 1, 2, 3*, 4, 5, 6, 7, 8, 9, 10, 11*, 12*, 14, 15*, 16, 17, 18, 19, 20, 21, 22, 31, 33, 34, 62, 69, 72, 75, 81, 86, 107, 108, 117*, 118*, 120*, 121*, 122*, 123*, 124 acrophylla 94 acrophylla var. ulei 94 acutata 47 acutiflora 9, 13, 17, 21, 28, 86-88, 89*, 93, 99, 105, 123* alpigena 12, 13, 27, 75, 76-81, 82* alpigena var. alpigena 9, 17, 21, 25, 26, 28, 76, 77, 78-79, 80, 82*, 86, 122* alpigena var. longifolia 9, 21, 26, 77, 78, 81, 82* alpigena var. rufa 9, 15*, 16, 17, 21, 24, 25, 28, 29, 74, 75, 76, 77, 78, 79-81, 82*, 83, 122* subsect. Aovatae 20 apiculata 17 bracteosa 9, 13, 19, 21, 25, 27, 28, 78, 80, 81-83, 86, 87* bracteosa var. alpigena 78 bracteosa var. australis 79

134 bracteosa var. expallens 82 bracteosa var. franciscensis 75 bracteosa var. guaratubensis 78 bracteosa var. ibitipocensis 82 bracteosa var. seriato-pedunculata 79 brevipedicellata 9, 13, 19, 21, 26, 27, 102, 104, 106*, 107, 123* bridgesii 112 buxifolia 62 cambessedeana 81, 82, 83 campestris 9, 13, 16, 17, 21, 23, 24, 31, 33-34, 41*, 117* campestris var. distans 33 camphorata 35 candolleana 107 catharinae 78 chilensis 49 chrysocarpa 9, 13, 18, 21, 25, 30, 57, 62-63, 66, 69*, 71, 121* colchaguensis 9, 13, 21, 125, 74-75, 78, 80, 82*, 122* correifolia 9, 13, 16, 21, 29, 57, 60*, 61-62, 71, 75, 78 cucullata 9, 13, 22, 26, 27, 28, 29, 47, 70*, 72-73, 121* subsect. Dichotomae 20 diemii 108, 111 distans 33 distoma 56 estrellensis 72, 73, 94 euosma 9, 12*, 13, 17, 19, 22, 24, 25, 27, 28, 69, 76, 78, 79, 83-86, 87*, 113, 114, 115, 116 euosma x glaucescens 113-116 euosma var. oblongata 84 exsucca 9, 13, 15*, 19, 21, 23, 24, 35-37, 39, 41*, 44, 108, 109, 110, 111, 112, 113, 117* exsucca var. bridgesii 112 exsucca x lanceolata 112-113 exsucca x ovata var. nannophylla 108-112 exsucca var. quetrihuensis 108 fernandeziana 9, 10, 13, 14, 17, 20, 21, 22, 23, 24, 30-31, 33, 41* ferreira-limana 94 ferruginea 40 filibracteata 103 franciscensis 13, 21, 28, 75-76, 78, 87*, 122* fuscovelutina 96, 99 glaucescens 13, 16, 25, 30, 39, 42, 47, 63-66, 69, 71, 82, 86, 113, 114, 115, 116 glaucescens forma catharinensis 67 glaucescens forma debilis 67 glaucescens forma pallida 67 glaucescens var. glaucescens 9, 22, 64, 65, 66-9*, 71, 113, 114, 115, 116, 121* glaucescens var. latior 9, 17, 21, 23, 47, 64, 65, 66, 67, 69*-71, grisea 67 hatschbachii 9, 13, 19, 21, 26, 39, 46, 48, 52-54, 55*, 56, 119* hoehnei 9, 13, 21, 24, 33, 103-104, 106* subsect. Incanae 20 itatiaiensis 45

Flora Neotropica johowi 61 jonssonii 32 kleinii 9, 13, 21, 23, 24, 29, 57, 58, 59, 60*, 62, 71, 120* lanceolata 9, 12*, 13, 17, 21, 24, 26, 27, 37-40, 41*, 48, 53, 112, 113, 117* latior 70 lechleriana 35 leptocalyx 9, 13, 22, 28, 102, 104, 106*-107 leptocalyx var. coriacea 106 leptorhyncha 78 leptospermoides 9, 13, 21, 27, 48, 51-52, 55*, 56, 120* longifolia 81 longipedicellata 107 longipedunculata 75, 122* luma 30, 31 sect. Macrobothrys 20 macrocalyx var. obovata 105 macrosepala 20, 105 malvillana 74 maritima 61 sect. Microbothrys 20 miersiana 9, 10, 13, 17, 19, 21, 24, 25, 26, 27, 28, 33, 90, 93, 96, 97, 99-100, 101*, 103 miersiana var. lanceolata 88, 90, 100 miersiana var. macahensis 96, 97 miersiana var. venosa 96, 99 montana 44, 81 montevideensis 101 monticola 82, 83 mosenii 60, 107 mucronata 75 multiflora 35 myrcioides 13, 14, 23, 73, 88, 89*, 90, 91-95, 103 myrcioides var. acrophylla 9, 15*, 17, 21, 26, 73, 89*, 91, 92, 94-95 myrcioides var. hypericifolia 92 myrcioides var. loefgreniana 92 myrcioides var. myrcioides 9, 21, 88, 89*, 91, 92-94, 123* myrcioides var. paranensis 94, 95 myrcioides var. ulei 94, 95 myrtoides 9, 13, 22, 24, 25, 100-101*-103, 123* myrtoides var. conferta 100, 102 myrtoides var. myrtoides 100 myrtoides var. stricta 100, 102 nannophylla 44 nothorufa 96, 98 nothorufa var. major 96, 98 nothorufa var. venosa 96, 99 obtusa 9, 12*, 13, 21, 24, 29, 49-50, 55*, 75, 118* obtusa var. berteroana 49 obtusa var. polyantha 49 obtusa var. raran 49 ovata 13, 14, 29, 42-47, 48, 55*, 64, 65, 66, 67, 114 ovata var. acutata 9, 17, 21, 23, 25, 29, 43, 47, 55*, 64, 65, 66, 67, 71, 73 ovata var. gracilis 9, 16, 17, 19, 21, 26, 43,

135

Myrceugenia 45-47, 51, 53, 54, 55*, 56, 64, 65, 66, 67, 114 ovata var. nannophylla 9, 21, 37, 42, 43, 44-45, 48, 55*, 66, 108, 109, 110, 111, 112 ovata var. ovata 9, 21, 42, 43-44, 55*, 66, 118* oxysepala 9, 12*, 13, 15*, 17, 20, 21, 27, 102, 104, 105-106*, 107 pallida 67 parvifolia 9, 13, 21, 26, 48, 50-51, 52, 55*, 118* sect. Phanerocalyx 20 pinnifolia 9, 13, 21, 27, 39, 45, 47-49, 53, 55*, 66 pilotantha 13, 24, 26, 28, 33, 93, 95-98*-99, 100, 103 pilotantha var. major 9, 15*, 19, 22, 88, 96, 97, 98*-99 pilotantha var. pilotantha 9, 17, 21, 96, 9798* planipes 9, 13, 15*, 16, 17, 21, 23, 27, 29, 56-58, 60*, 62, 63, 66, 71, 120* ramboi 96, 98 regnelliana 45, 46, 114 regnelliana var. capillaris 45 regnelliana var. coriacea 45 regnelliana var. var. dubia 113, 114 regnelliana var. gracilis 45 regnelliana forma itatiaiensis 45 regnelliana var. leptophylla 45, 54 reitzii 9, 13, 16, 17, 22, 23, 57, 58-60*, 62, 71, 107, 120* rufa 9, 13, 16, 21, 25, 40-41*-42, 75, 118* rufescens 9, 10, 13, 15*, 16, 17, 21, 23, 24, 25, 31, 32-33, 41*, 97, 98, 103, 104, 117* rufescens var. alegrensis 32 schultzei 9, 12*, 13, 17, 21, 24, 25, 34-35, 41* scutellata 9, 13, 21, 25, 70*, 71-72 sellowiana 101, 107 seriatoramosa 9, 13, 21, 26, 27, 28, 29, 70*, 73-74, 78, 80, 121* smithii 9, 13, 19, 22, 27, 46, 47, 48, 52, 53, 54-55*-56, 119* sect. Steganocalyx 20 stenophylla 38 stenophylla var. pinifolia 47 subsect. Subulatae 20 thalassaia 61 subsect. Uniflorae 20 valientei 43, 44 venosa 9, 13, 22, 28, 88-89*-90, 93, 94, 100 Myrcia 2, 5, 6, 14, 16, 19 subgen. myrceugenia 20 planipes 56

Myrcianthes 5, 7 fragrans 19 Myrciaria 5 Myrciinae 2, 4*, 5, 6, 7, 14, 16, 19 Myrcioideae 5 Myrtaceae 1, 2, 3, 4, 5, 7, 13, 14, 17, 18, 19, 124 Myrteae 1, 2, 4*, 5, 6, 7, 10, 18, 19 Myrteola 5, 18 Myrtinae 4*, 5, 6, 7 Myrtoideae 4 Myrtus 2, 5 brachyphylla 51 communis 17, 18 femandeziana 30 gudilla 38 lanceolata 37, 38 luma 31 maxima 31 raran 49 rufa 40 Neomyrtus 18 pedunculata 18 Nothofagus antarctica 44 dombeyi 63 Nothomyrcia 6, 20 fernandeziana 20, 30, 31 maxima 31 Paivaea 5 Pimenta 5, 31 Pimentoideae 5 Plinia 5 Pseudanamomis 5 Pseudocaryophyllus 5 Psidium 5, 18, 107 incanum 107 itatiaiae 78 Reichea 5, 7 Siphoneugena 5 Sophora 113 Temu 5, 6, 7, 19 Tepualia 4 Thryptomene 18 Ugni 5, 18 Verticordia 18