rt 7+ yi PHYTOLOGIA eo An international journal to expedite plant systematic, phytogeographical and ecological publication Vol. 68 February 1990 No.2
CONTENTS
“4 M. CHEEK, A new species of pygmy Drosera from Western Australia and a note on the status of sect. Bryastrum and sect. I eta cl aa ay gacaagyeocannderasibsckeess danse’ 85
y B.L. TURNER, Two new species of Brickellia (Asteraceae: Eupatorieae) from northern MEXICO o.oo... eeeeceeeecceeesseeeeeeneeeeeeees 90
B.L. TURNER, Ageratina moorei (Asteraceae: Eupatorieae), a new species from the state of MExicO, MEXICO oe ccc eeeeeseceeeeeeeees 95
B.L. TURNER, Two new species of Verbesina (Asteraceae: eT eT oa ic. 2 hyo ein aR eg an SS 98
V. RICO-GRAY, Aizoaceae de la Peninsula de Yucatan, México . 102 ~ V. RICO-GRAY, Bataceae de la Peninsula de Yucatan, México ... 108
V. RICO-GRAY, Rhizophoraceae de la Peninsula de Yucatan, Pee Este ek ae dO ta RO a a a RON 113
~ A.C. NICOLSON, A new species of Miconia (Melastomataceae) cccerpeg toate) FU at iy Bo ca 4 0)”: sa PN RL oe mn 120
Contents continued on the inside cover.
Published by Michael J. Warnock 185 Westridge Drive Huntsville, Texas 77340 U.S.A. PHYTOLOGIA is printed on acid free paper.
Price of this number $3.50; for this volume $20.00 to institutions in advance or $18.00 to individuals in advance; back volume prices apply if payment is received after a vulume is closed; 512 pages constitute a complete volume, claims for numbers lost in the mail must be made immediately after receipt of the next following number for free replacement.
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“- PHYTOLOGIA
An international journal to expedite plant systematic, phytogeographical and ecological publication
Vol. 68 February 1990 No.2
CONTENTS
’ M. CHEEK, A new species of pygmy Drosera from Western Australia and a note on the status of sect. Bryastrum and sect. eign nas cpl Bontek row ee Raw Aes waiePrarahinededeean bee iaei: 85
- B.L. TURNER, Two new species of Brickellia (Asteraceae: Eupatorieae) from northern MEXICO .0....... cee eccsesteceseneeeesneeeeenees 90
B.L. TURNER, Ageratina moorei (Asteraceae: Eupatorieae), a new species from the state Of MExicO, MEXICO oe... eeeceeeeeeeeeees 95
B.L. TURNER, Two new species of Verbesina (Asteraceae: Reet GY TENT) MOKICO. 20 coc sce sc one ssceetonee cece 98
V. RICO-GRAY, Aizoaceae de la Peninsula de Yucatan, México . 102 ~ V. RICO-GRAY, Bataceae de la Peninsula de Yucatan, México ... 108
V. RICO-GRAY, Rhizophoraceae de la Peninsula de Yucatan, (ONT te Sich 2M Sg ig a RP LSE, Ba eee, rh 113
. A.C. NICOLSON, A new species of Miconia (Melastomataceae) Benir erate. 1 OSSCr ANTS oko ioic ccesactacess Lane sceiona teense 120
Contents continued on the inside cover.
Published by Michael J. Warnock 185 Westridge Drive Huntsville, Texas 77340 U.S.A. PHYTOLOGIA is printed on acid free paper.
Price of this number $3.50; for this volume $20.00 to institutions in advance or $18.00 to individuals in advance; back volume prices apply if payment is received after a vulume is closed; 512 pages constitute a complete volume, claims for numbers lost in the mail must be made immediately after receipt of the next following number for free replacement.
A
_G.L. NESOM, Two new species of Archibaccharis (Asteraceae: : Astereae) from Oaxaca, MEXICO ...........c<seccesescssecconcsnense ee 122°
E.E. TERRELL, Synopsis of O/den/andia (Rubiaceae) in the United ri no ea 1a
B.L. TURNER, New names and combinations in Mexican Alloispermum (Asteraceae: Heliantheae) ..............cccccceeeeee 134~
J. OLSEN, Corrections in the type locality citation of Verbesina _ i Lo Cee ae eens, EMRE Ie 137
B.L. TURNER, A new species of Aphanactis (Asteraceae: Heliantheae; Galinsoginae), from Oaxaca, México .............. 138
P.M. MCKENZIE, L.E. URBATSCH & M.G. LELONG, Panicum rigidulum var. combsii (Poaceae) in Louisiana ..................000 141
G.L. NESOM, Taxonomic summary of Ericameria (Asteraceae: Astereae), with the inclusion of Haplopappus sects. MACTOREGHTTA ANG ASHES. oo... sc0cc0cecss00cceesnceneeovseentcesasise en 144
RCO NS PRC oir oso icceiecewccdcccnecce ctncasscnedencucnustaveedceederee eee 156
i. nana p AO. 7 BOTANICAL GARDEN
Phytologia (February 1990) 68(2):85-89.
A NEW SPECIES OF PYGMY DROSERA FROM WESTERN AUSTRALIA AND A NOTE ON THE STATUS OF SECT. BRYASTRUM AND SECT. LAMPROLEPIS
Martin Cheek Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K.
ABSTRACT
Drosera mannii Cheek, a species widely known in cultivation as Drosera ‘Bannister’ since 1977 or earlier, is described. This opportu- nity is taken to re-evaluate the widely maintained distinction between Drosera sect. Bryastrum Planch. and Drosera sect. Lamprolepis Planch.
KEY WORDS: Drosera, Droseraceae, taxonomy, Australia.
The preparation of accounts of the cultivated species of Drosera for the European Garden Flora Project and the Royal Horticultural Society’s new Dictionary of Gardening (both in prep.) prompted a critical examination of several taxa of pygmy Drosera of West Australian origin which lack species epithets and are known solely by cultivar names based on the locality from which they were collected. Many of these have been widely cultivated in Europe and North America since the late 1970’s. The first time that these were discussed was apparently by Rose (1977), who mentioned 9 such taxa: ‘Toodyay Pink,’ ‘Mt. Manypeaks,’ ‘Muchea Pink,’ ‘Regan’s Ford,’ ‘Lake Bad- gerup’ (misprinted as ‘Lake Badgebup’), ‘Bannister,’ ‘Walyinga,’ ‘Brookton’ and ‘North Beermullah.’ Slack (1979: 219) expounds upon the cultivation of many of these and others (‘Millbrook Road’ and ‘Moora’), and it is clear that even then, several were well known in cultivation in Europe. Two addi- tional cultivars, ‘Gidgeganup Pink’ and ‘Gidgeganup White’ were discussed by Lowrie (1980) and their introduction attributed to Rose. Although all these taxa seem to have been considered as putative new species by Rose (1977), in- vestigation has shown that this is not so for all of them. Drosera ‘Gidgeganup Pink’ for example, is clearly only a pink strain of Drosera scorpiotdes Planch. and should be cited as D. scorpioides ‘Gidgeganup Pink.’ However, some of these taxa do seem to represent distinct species which are unaccounted for in the recent treatment of the genus in the Flora of Australia (Marchant, et al. 1982). Subsequently, one of these taxa (‘Regan’s Ford’) has been described by Strid (1987) as D. rechingeri, but the bulk of the taxa are still botanically
85
86 PHYTOLOGIA volume 68(2):85-89 February 1990
unaccounted for. Although it is not possible, owing to lack of materials, to properly assess all of these taxa, it is plain that at least one of those still in widespread cultivation in Europe and America, described by both Rose (1977) and lengthily by Slack (1979) is undoubtedly specifically distinct. Accordingly, it is described as new here.
Drosera mannii Cheek, sp. nov. TYPUS: AUSTRALIA. West Australia: Cheek 2064 (HOLOTYPUS: K; Isotypi: PERTH,CANB,P).
Drosera ‘Bannister’ S. Rose, Carn. Pl. Newsl. 6(1):11. 1977.
A Drosera leucoblasta Bentham stylis 5 vel 6, stigmatibus sub- obclavatis, petalorum unguibus 1 mm et laminis retusimissis dif- fert.
Herbaceous, carnivorous, rosulate perennial. Stem short, unbranched, erect, base surrounded by marcescent leaves and stipules. Roots 3-7, wiry, largely adventitious, 0.2 mm thick and ca 5 cm long, dull red or orange above, straw coloured below with 5-11 lateral branches and numerous black, fine hairs ca 1.5 mm long. Leaves numerous, leaf blade elliptic, ca 2.5 mm long, 2 mm wide, concave in T.S., distal glandular hairs ca 4 mm long, proximal hairs ca 2 mm long, underside glabrous; petiole 3.5-4 mm long, 0.5-1 mm broad in plane view, dorsiventrally flattened, glabrous; stipules not detaching eas- ily with the petioles from the stem, 3.5-4 mm long, 1 mm wide at the base, dividing ca 1.5 mm from the base into 3 main teeth, each spreading, trian- gular, 2 mm long with a laciniate margin, membranous-chartaceous, hyaline. Inflorescences 1-2, the sterile part 7-10 cm long, sub-glabrous, with extremely few, scattered, randomly directed, appressed white hairs, the upper 5-8 mm glandular hairy, the whole flushed dark red; fertile part 3-8.5 cm long, bearing up to 32 flowers, glandular hairy. Flowers 12-15 mm across, ascending. Sepals 5, ovate-elliptic, 2.5 mm long, 2 mm broad, acute-obtuse, very finely toothed, outer surface with red glandular hairs. Petals 5(-6-7), imbricate; petal blade oblong-obovate, 5-7(-8) mm long, 4.5-6 mm wide, apical notch 1 mm deep, the base truncate, palest pink (R.H.S. colour chart ‘red-purple group’ 62D), with a very thin red margin; petal stalk ca 1 mm long, broadest at tip, green. Sta- mens 5(-6) ascending; filaments 2.5-3 mm long, 0.25 mm wide at apex (which confluent with the connective), tapering to the base, white; anthers ca 0.3 mm long, with 2 separate thecae, inserted obliquely on either side of the swollen connective, purple; pollen orange. Ovary ca 0.75 mm diam, white. Styles 5-6(-7), 2 mm long, filiform. Stigma obconical to obclavate, 1.25 mm long, 0.5-0.75 mm wide, translucent white, style insertion ventral and oblique, 0-5 mm from base. Fruits with pedicels recurved, seeds unknown.
WESTERN AUSTRALIA: ez cult., 1984 Adrian Slack from plants origi- nally found by Phillip Mann at the side of the Bannister road, southwestern W. Australia, Cheek 2064 (HOLOTYPE: K; Isotypes: PERTH,CANB,P).
Cheek:
New Drosera from Western Australia
Figure 1. Drosera mannti. A. habit; B. sepal; C. clawed, retuse petal; D. infructescence with bract; E. peltate, obclavate stigma; F. stamens; G. flower in plane view; H. stipule; I. leaf. All drawn from Cheek 2064 by the author. Scale bar = 1 mm.
87
88 PHYTOLOGIA volume 68(2):85-89 February 1990
TABLE 1: Characters distinguishing D. mannit, D. leucoblasta and D. pul- chella.
| D. mannii | D. leucoblasta | D. pulchella
1-0.75 mm | <0.5-0.75 mm | 1-2.25 mm
Petiole breadth
Petal apex retuse entire entire Stigma shape obconical filiform filiform to -obclavate clavate Stigma number 5-6 3-5 4-5 Fruits descending | descending ascending
HABITAT. In pine leaf mould on the edge of pine forests, on damp creek- sides and in wet hollows (Rose 1977).
The curious and absolutely distinctive stigmas that characterize this species were first and aptly described (as ‘boat-shaped’) by Rose (1977), in his dis- cussion of D. ‘Bannister.’ A long description was given by Slack (1979) who placed it near D. pulchella Lehm. Although it shares with that species un- usually broad petioles and pink flowers, it is clearly more closely related to D. leucoblasta Benth. Indeed, in the key of Marchant, et al. (1982), it keys out to couplet 70, but falls midway between the two opposing statements, having styles 2 mm long, scape 9-15 cm. It is readily distinguished from D. leucoblasta Benth. by its stigmas (probably the most important single source of specific and sectional characters in the genus) and emarginate petals, and from D. pulchella Lehm. which also has comparatively wide petioles and occasionally pink flowers, by these same characters. The differences between these taxa are elucidated further in Table 1.
The species is widely cultivated, being readily propagated, predominantly from the small, disc like gemmae produced in December (Northern Hemi- sphere). A single inflorescence can produce a succession of flowers from June to early September.
Sectional delimitation
Planchon’s sectional classification of Drosera divided the perennial, gem- mae forming pygmy sundews into two groups: sect. Lamprolepis Planch., with 8 species characterized by 5 stamens, 3-5 styles, inflorescences many flow- ered and leaves rarely peltate, and sect. Bryastrum with a single species, D. pygmaea DC., characterized by tetramerous flowers, 4 styles, inflorescences single flowered, leaves peltate. D. pygmaea was maintained as distinct from section Lamprolepis by Marchant (1982), but in view of the nature of D. occt- dentalis Morrison (1912), D. ‘Lake Badgerup’ and D. ‘Warriup,’ the value of maintaining this distinction seems dubious. D. occidentalis has 1-2 flowered inflorescences with pentamerous flowers having 4-5 styles. D. ‘Lake Badgerup’
Cheek: New Drosera from Western Australia 89
usually has 1 flowered inflorescences, but occasionally produces up to 7 flowers per scape, the flowers being 4-5-merous, with 4 or 5 styles. D. ‘Warriup’ is uniflowered, is pentamerous, with 3 styles. While it is true that all 3 have peltate leaves, this character also occurs in species of section Lamprolepis, for example, D. pulchella Lehm. In short, on present evidence, it seems best to place D. pygmaea in the same section as the remainder of the pygmy sundews.
ACKNOWLEDGMENTS
I am very grateful to Melanie Thomas for help with the diagnosis and to David Hunt for comments on the manuscript.
LITERATURE CITED
Diels, L. 1906. Droseraceae in A. Engler (Ed.), Das Pflanzenreich 26: 1-136.
Lowrie, A. 1980. Another look at Drosera species. Carn. Pl. Newsl. Austral. 5:4.
Marchant, N.G., H.I. Aston & A.S. George. 1982. Droseraceae in Flora of Australia 8:9-66.
Morrison, A. 1912. New and rare Australian plants. J. Bot. 50:164-168.
Planchon, J.E. 1848. Sur la famille des Droséracées. Ann. Sci. Nat. Bot. 9:79-99.
Planchon, J.E. 1848. Sur la famille des Droséracées. Ann. Sci. Nat. Bot. 9:185-207.
Planchon, J.E. 1848. Sur la famille des Droséracées. Ann. Sci. Nat. Bot. 9:285-309.
Rose, S. 1977. Seeking the pygmy droseras. Carn. Pl. Newsl. 6(1):10-11 and figs. 10.1-10.4.
Slack, A. 1979. Carnivorous Plants. Ebury Press, London (138-141 and 219).
Strid, A. 1987. New species of Beaufortia and Chamaelaucium (Myrtaceae), Drosera (Droseraceae) and Pultenaea (Fabaceae) from S.W. Australia.
Pl. Syst. Evol. 155:343-345.
Phytologia (February 1990) 68(2):90-94.
TWO NEW SPECIES OF BRICKELLIA (ASTERACEAE: EUPATORIEAE) FROM NORTHERN MEXICO
B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Brickellia wendtii B. Turner, from northern Coahuila and B. worthingtonii, from southern Durango, are newly described and illus- trated. The former, with nonplumose pappus bristles, is closely related to B. secundiflora (Lag.) A. Gray. The latter, with subplumose pappus bristles, is closely related to B. oretthales (B.L. Robins.) Shinners.
KEY WORDS: Brickellia, Asteraceae, Eupatorieae, taxonomy, neotrop- ical flora, México.
Preparation of a treatment of the genus Brickellia for México has necessi- tated description of the following two novelties.
Brickellia wendtii B. Turner, sp. nov., Figure 1. TYPE: MEXICO. Coahuila: Mpio. de Muzquiz, Rincon de Maria (28° 27’ 30” N, 102° 04’ W), “in roadbed at spring level in deciduous woodland,” ca 1400 m, 23 Aug 1975, T. Wendt, E. Lott & D.H. Riskind 1264 (HOLOTYPE: TEX; Isotype: MEXU).
Brickellia secundiflorae (Lag.) A. Gray similis sed capitulis flosculis 10-14 in cyma terminali ramosa dispositis, pedunculis ul- timis erectis 2-3 mm longis, receptaculis glabris, et acheniis 6.0-6.5 mm longis differt.
Many stemmed suffruticose perennial herbs to 1 m high. Stems densely glandular pubescent, mostly simple and arising from a woody rootstock. Leaves mostly opposite, 5-8 cm long, 2.5-4.0 cm wide, glandular pubescent; blades ovate, sparsely rough hispid above and below, glabrate with age, trinervate, the secondary veins weakly developed, the margins crenuloserrate; petioles 3- 12 mm long. Heads ca 15 in a terminal open cyme, the ultimate peduncles mostly 2-3 cm long, densely glandular pubescent, erect or ascending. Involu- cres 12-13 mm high, turbocampanulate, the bracts 3-4 seriate, the apices acute or apiculate. Receptacle glabrous. Florets 11-12 per head; corollas ca 7 mm long, reportedly “creme green”. Achenes 6.0-6.5 mm long, sparsely pubescent
90
New species of Brickellia from Mexico 91
Turner:
Fig.1. Brickellia wendtii, from holotype.
92 PPOs OL OSTA volume 68(2):90-94 February 1990
with appressed hairs, the pappus of ca 40 white barbellate bristles ca 6 mm long.
The species is superficially similar to B. secundzflora but the heads are wholly terminal and nonpendulous, the florets fewer per head, the receptacle glabrous and the achenes longer. In Robinson’s (1917) treatment, the species will key to B. saltillensis B.L. Robins., which I have treated as synonymous with B. secundiflora (Turner 1989).
It is a pleasure to name this distinct taxon for my friend and oft-time col- league, Dr. T. Wendt, currently working on the Mexican flora out of Chapingo, Mexico (CHAPA).
Brickellia worthingtonii B. Turner, sp. nov., Figure 2. TYPE: MEXICO. Durango: ca 75 km S of Cd. Durango, highway to La Flor, 6 km by road from La Flor on the road to Cd. Durango (ca 23° 33’ N, 104° 42' W), 8500-9000 ft, pine forests on rocky slopes, 18 Aug 1982, R.D. Worthington 8838 (HOLOTYPE: TEX; Isotype: MEXU).
Brickellia oretthales (B.L. Robins.) Shinners similis sed capit- ulis majoribus paucioribus in pedunculis ultimis longioribus et foliis ovatis 2-3-plo longioribus quam latioribus differt.
Slender perennial herbs 30-60 cm high. Stems simple, mostly unbranched, minutely scabrid hispidulous, arising from slender rhizomes. Leaves mostly alternate (only those at a few lower nodes opposite), 20-40 mm long, 8-12 mm wide; petioles 1-3 mm long; blades ovate to ovate elliptic, sparsely minutely scabridulous, especially on the nerves and along the margins, denticulate to nearly entire. Heads 6-10 per stem, arranged in simple secund racemes, the ultimate peduncles 1-4 cm long, drooping. Involucres narrowly campanulate, 12-14 mm long, the bracts ca tetraseriate, glabrate with puberulous-ciliate margins. Receptacle glabrous. Florets 10-11 per head; corollas 8-9 mm long. Achenes ca 4 mm long, densely sericeous throughout, the pappus of ca 35 white subplumose bristles 8-9 mm long.
Additional collection examined: México. Durango: Mpio. de Suchil, arroyo El] Taray, Reserva de la Michilia, pine forest along arroyos, 20 Sep 1985, J. Alvarado 44 (CIIDIR,TEX).
Because of its secund, drooping heads, the species superficially resembles Brickellia secundiflora. It is actually more closely related to B. oreithales, possessing the subplumose pappus and drooping heads of the latter, but differs markedly by its fewer larger heads and ovate leaves.
It is a pleasure to name this taxon for Dr. R.D. Worthington, Professor in the Department of Biology, University of Texas, El Paso, whose explorations in northern México have added several plant novelties to the region.
93
New species of Brickellia from México
Turner:
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— SSS ee ~ ay —_—_—_— =s e: ‘en ase ——_ mae <
from holotype.
gtoni,
Brickellia worthin
Rag. 2:
94 PHYTOLOGIA volume 68(2):90-94 February 1990
ACKNOWLEDGMENTS
I am grateful to Dr. Guy Nesom for the Latin diagnoses and to both him and Dr. Carol Todzia for a review of the manuscript itself. Nancy Webber provided the illustrations.
LITERATURE CITED
Robinson, B.L. 1917. A monograph of the genus Brickellia. Mem. Gray Herb. 1:1-151.
Turner, B.L. 1989. Conspectus of the Mexican species, Brickellia secundiflora (Lag.) A. Gray, and its infraspecific categories. Phytologia 67:307-311.
Phytologia (February 1990) 68(2):95-97.
AGERATINA MOOREI (ASTERACEAE: EUPATORIEAE), A NEW SPECIES FROM THE STATE OF MEXICO, MEXICO
B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Ageratina moorei B. Turner, is described and illustrated. It is known from only two collections, both obtained in or near Valle del Bravo, western portion of the state of México. It is closely related to A. cardiophylla (B.L. Robins.) King & H. Robins., but is readily distinguished by its smaller habit, mostly basal leaves and smaller heads with fewer florets.
KEY WORDS: Ageratina, Asteraceae, Eupatorieae, taxonomy, Mex- ican flora.
Routine identification of Mexican Asteraceae has revealed the following novelty.
Ageratina moorei B. Turner, sp. nov., Figure 1. TYPE: MEXICO. México: Mpio. Valle del Bravo, moist ravine by Rio Molino in pine-oak woods above reservoir, 1-2 Nov 1949, H.E. Moore, Jr. & Mar Cetto 5470 (HOLOTYPE: BH; Isotype: BH).
Ageratina cardiophyllae (B.L. Robins.) King & H. Robins. sim- ilis sed plantis minoribus (50-60 cm altis), foliis plerumque basal- ibus, et involucris brevioribus (5-6 mm vs 7-9 mm) differt.
Perennial herbs 50-60 cm high. Leaves opposite, mostly basal, the upper ones remote and much reduced; petioles 3-9 cm long, pubescent with crinkly spreading hairs; blades neatly cordate, mostly 5-13 cm long, 4-9 cm wide, 3-5 nervate from the base, moderately pubescent above and below, especially along the major veins, the margins rather evenly crenulate; heads numerous in rather open terminal cymes, the ultimate peduncles mostly 8-15 mm long, densely pubescent with glandular trichomes. Involucres turbocampanulate, mostly 5-6 mm high, the bracts glandular pubescent with acute apices. Receptacle plane, glabrous. Florets 13-16 per head, the corollas pinkish white, glabrous, ca 4
95
96 PHYTOLOGIA volume 68(2):95-97 February 1990
Fig.1. Ageratina moorei, from holotype.
Turner: New species of Ageratina from México 97
mm long. Achenes 2.0-2.5 mm long, hispidulous; pappus of ca 20 fragile barbellate bristles 4-5 mm long in a single series.
Additional specimen examined: México. México: 15 mi E of Temascaltepec along highway 134, N facing slopes, S of highway, oak-pine woods, 28 Oct 1981, M.J. Warnock 2591 (TEX).
The species is closely related to A. cardtophylla and A. macvaughu King & H. Robins., but can be distinguished from both by its smaller habit with leaves mostly basal. It is perhaps closest to A. macvaughit, possessing the small heads and florets of that species, but differs in habit, as noted, and by its neatly cordate leaves which are 3-5 palmately nerved from the very base, whilst those of A. macvaughi are 3-5 subpalmately nervate from above the base.
It is a pleasure to name this species for its first collector, the late Professor H.E. Moore, Jr., working out of the L.H. Bailey Hortorium, Cornell University. He assembled a rich set of collections from México over a 20 year period (mostly
1940-60).
ACKNOWLEDGMENTS
I am grateful to Dr. Guy Nesom for the Latin diagnosis and to him and Dr. T. Wendt for reviewing the manuscript itself. Ms. Nancy Webber provided the illustration.
Phytologia (February 1990) 68(2):98-101.
TWO NEW SPECIES OF VERBESINA (ASTERACEAE: HELIANTHEAE) FROM MEXICO
B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Two new species of Verbesina, V. hidalgoana B. Turner and V. macdonaldii B. Turner are described from Hidalgo and Oaxaca, México, respectively. Verbesina hidalgoana belongs to the section Pseudomon- tanoa; the relationship of V. macdonaldit is obscure, apparently repre- senting a very distinct element in the genus.
KEY WORDS: Verbesina, Asteraceae, Heliantheae, Hidalgo, Oax-
aca, taxonomy, México.
Routine identification of Mexican Asteraceae has revealed the following novelties.
Verbesina hidalgoana B. Turner, sp. nov. TYPE: MEXICO. Hidalgo: Mpio. Molango, vicinity of Molango on road to Lolotla, 1600 m, 9 Nov 1946, H.E. Moore, Jr. 1979 (HOLOTYPE: BH).
Verbesina fastigiatae B.L. Robins. & Greenm. multo similis sed foliis paucioribus omnino nonlobatis et capitulescentia multo pau- ciore foliis superis tantum parum longiore differt.
Suffruticose herbs or shrubs 1-2 m high. Stems hispidulous, narrowly winged throughout, the wings 0.5-1.0 mm wide. Leaves opposite through- out, 10-22 cm long, 5-10 cm wide; petioles 2-7 cm long; blades ovate-deltoid, gradually tapering upon the petioles, the latter winged throughout, rather coarsely pubescent above with broad based, recurved hairs, the undersurfaces coarsely hispidulous with straight hairs, especially along the venation, the mar- gins serrulate, unlobed. Heads 20-50 in terminal congested cymose panicles, the ultimate peduncles mostly 3-15 mm long, the capitulescence 4-8 cm wide, 4-7 cm high, only slightly exceeding the upper foliage. Involucres turbocam- panulate, 4-6 mm high, the bracts graduate, hispidulous. Ray florets 8-13, pistillate, fertile, the ligules yellow, 4-6 mm long. Disk florets ca 30; corollas yellow, the tube pubescent, ca 1 mm long, the throat glabrous, ca 3 mm long.
98
Turner: New species of Verbesina from México 99
Achenes 4-5 mm long, glabrous or nearly so, the pappus of 2 awns 3-5 mm long.
Additional specimen examined: MEXICO. Hidalgo: Mpio Zacualtipan, vicinity of Zacualtipan, rich mixed woods in ravine beside Rio Teponapa, ca 2000 m, 20 Oct 1949, H.E. Moore, Jr. 5389 (BH).
Verbesina hidalgoana clearly belongs to the section Pseudomontanoa (Turner 1985) where it relates to V. fastigzata and V. olseniz B. Turner, differing from both in its smaller unlobed leaves and reduced capitulescence which barely exceeds the upper foliage.
Verbesina macdonaldii B. Turner, sp. nov., Figure 1. TYPE: MEXICO. Oaxaca: 35 km ESE of Miahuatlan, 5 km NE of Santo Domingo Ozolote- pec, Cerro Quiexobra, timberline vegetation in open glades along ridges and in mountain “saddles,” occasional on SE exposures, 3650-3800 m, (16° 10’ N, 96° 15’ W), 10 Dec 1989, Andrew McDonald 2936 (HOLO- TYPE: TEX; Isotype: MEXU).
Verbesina oazacanae DC. similis sed foliis oppositis crassioribus in paginis infernis dense appressi-pilosis et capitulis paucioribus involucris ca 1.5 cm altis differt.
Woody shrubs 2-3 m high. Stems densely pubescent, wingless. Leaves opposite, thick and leathery, 8-12 cm long, 3-4 cm wide; petioles thick, 2- 4 mm long; blades mostly elliptical, somewhat bicolored, moderately hirsute and dark green above, densely pilose and dirty white beneath, the margin rev- olute, entire or nearly so. Heads radiate, arranged 3 to 4 in terminal cymes, the ultimate peduncles 3-6 cm long, thick and densely pilose. Involucres ca 1.5 cm high, ca 3 cm wide, 2-3 seriate, the outer bracts ovate-lanceolate, some- what spreading, longer than those of the inner series. Receptacles convex, ca 8 mm across, the pales linear-oblanceolate, acute at the apices. Ray florets 13-21, neuter or pistillate, fertile or not, the ligules dark yellow, 11-15 mm long, 2-3 mm wide, the tube pubescent, ca 3 mm long. Disk florets numerous, the corollas yellow, ca 4.5 mm long, the tubes pilose, ca 1.5 mm long, the lobes 5, ca 0.6 mm long. Anther sacs purplish. Style branches with acute appendages. Achenes (somewhat immature?) ca 4 mm long, black, radially flattened, glabrous, ca 1.5 mm wide, wingless or seemingly so, a very nar- row wing (ca 0.1 mm wide) along the upper shoulders; pappus of 2, readily deciduous, linear-lanceolate bristles 2.5-3.0 mm long.
A very distinct species, not readily related to yet others, but perhaps closest to Verbesina hebmannii Schultz-Bip. or V. oaracana DC., both of which have smaller, more numerous heads and alternate leaves.
100 PHY:T-:OLOGLIA volume 68(2):98-101 February 1990
Fig.1 Verbesina macdonaldii, from holotype.
Turner: New species of Verbesina from Mexico | 101
It is a pleasure to name this remarkably distinct species for its only known collector, Dr. Andrew McDonald, apparently the first professional systematist to ascend Cerro Quiexobra. Among his collection of Asteraceae from this first venture, seven new species were discovered (Nesom, in prep.; Turner, in prep.). Additional field work on this isolated mountain is certain to yield additional novelties.
ACKNOWLEDGMENTS
I am grateful to the Bailey Hortorium, Cornell University (BH), for the loan of materials. Dr. Guy Nesom provided the Latin diagnoses and both he and Dr. Andrew McDonald reviewed the manuscript. Ms. Nancy Webber provided the illustration.
LITERATURE CITED
Turner, B.L. 1985. Revision of Verbesina sect. Pseudomontanoa (Aster-
aceae). Pl. Syst. Evol. 150:237-262.
Phytologia (February 1990) 68(2):102-107.
AIZOACEAE DE LA PENINSULA DE YUCATAN, MEXICO
Victor Rico-Gray Centro de Ecologia, UNAM, A.P. 70-275, 04510 Mexico, D.F. MEXICO
RESUMEN
Se presenta la descripcion de la familia Aizoaceae para la Peninsula de Yucatan, México (Estados de Yucatan, Campeche y Quintana Roo). Se describe taxonomicamentea Sesuvium portulacastrum y a Trianthema portulacastrum; se presentan datos de su distribucion y ejemplares ex- aminados.
ABSTRACT
A description of Aizoaceae from the Yucatan Peninsula of México (states of Yucatan, Campeche and Quintana Roo) is presented. Taxo- nomic descriptions of Sesuvium portulacastrum and Trianthema portula- castrum, along with data about their distribution and a list of specimens examined are presented.
PALABRAS CLAVE (KEY WORDS): Aizoaceae, Sesuvium portu- lacastrum, Trianthema portulacastrum, Yucatan, Campeche, Quin- tana Roo, México.
INTRODUCCION
Los miembros familia Aizoaceae Rudolphi se caracterizan por ser hierbas o subarbustos anuales o perennes; con hojas suculentas; flores solitarias, acti- nomorficas y hermafroditas, tépalos y estambres mas o menos numerosos, ovario superior a inferior, plurilocular, ocasionalmente unilocular, numero de estilos igual al de léculos en el ovario, pocos a numerosos ovulos en cada loculo o solitarios y el fruto capsular, locular o circuncisil con sernillas pequenas y numerosas (Rico-Gray 1979; Rzedowski 1979).
Se considera que esta familia tuvo su origen y diferenciacion en el oeste del antiguo continente de Gondwana, entre los periodos Paleoceno y Eoceno (Raven & Axelrod 1974). Es una familia basicamente africana, pero se le encuentra bien representada en el Mediterraneo, en Australia y en las regiones secas de los tropicos y subtropicos de ambos hemisferios.
102
Rico-Gray: Aizoaceae de la Peninsula Yucatan, México 103
La familia Aizoaceae presenta alrededor de 11 géneros, dependiendo de los limites que se adopten, y aproximadamente 2,500 especies (Lundell & Lundell 1983; Standley & Steyermark 1946). En Norteamérica se encuentran cinco generos, dos de los cuales crecen silvestres en la Peninsula de Yucatan (Sosa, et al. 1985; Standley 1930).
En México tenemos la presencia del género Mesembryanthemum ( M. cristal- linum L., M. blandum Haworth, MM. chilense Mol., M. cordifolium L., M. nod- iflorum L.), originario del sur de Africa. Algunas especies de este género se utilizan como plantas de ornato, como es el caso de M. cordiflorum y M. cristallinum, conocidos en diferentes lugares como “siempre viva.” Estas es- pecies se caracterizan por presentar el caliz pentalobulado y numerosos pétalos de color rosa-rojo muy intenso y hojas ovadas o cordeadas; las flores son un poco mayores que en las especies de los géneros silvestres.
CLAVE PARA GENEROS DE AIZOACEAE DE YUCATAN
Clave para géneros de la Familia Aizoaceae presentes en la Peninsula de Yucatan:
1. Ovario con 3-5 léculos; semillas numerosas; estambres 5-numerosos; hojas de un mismo par mas o menos del mismo tamano, linear lanceoladas, PMeMiINUIAMey Yr. se tees Sees ie cate tes wate ae arenes oe Sesuvium
1’ Ovario con 1 6 2 léculos; semillas pocas; estambres 5-10; hojas de un mismo par de diferente tamano, eliptico-lanceoladas, las estipulas escar- RANORS bil. 2c hE ORE A OL Pe ere ees Trianthema
SESUVIUM L., Syst. Nat., ed. 10. 1058. 1759.
Plantas postradas o ascendentes, suculentas, glabras; varios tallos alternos originados de una corta raiz tipica, por lo general de color rosa-morados. Hojas opuestas, las del mismo par mas o menos de igual tamano, la lamina cilindrica a mas o menos plana, el margen entero, exestipuladas; peciolos dilatados. Flores axilares, sesiles a pediceladas, los cinco tépalos unidos en la base; es- tambres 5, libres y con insercion alterna a los tépalos o numerosos insertos en forma continua en el tubo floral, mas pequenos que los tépalos, el filamento filiforme o subulado, las anteras pequenas, comunmente amarillento-rojizas; estilos 2-5, lineares, erectos, el estigma longitudinal, papiloso; ovario semi- infero, 2-5 loculos, los 6vulos numerosos, anacampilotropos, la placentacion axilar. Capsula membranosa, envuelta por el perianto, circuncisil cerca del centro; semillas numerosas, pequenas, lustrosas y obscuras.
Género con cerca de ocho especies distribuidas en los tropicos y subtropicos de ambos hemisferios (Bogle 1970; Correll & Correll 1972; Lot-Helgueras & Rico-Gray 1978). En la Peninsula de Yucatan se presenta una especie (Sosa,
et al. 1985).
104 PHYTOLOGIA volume 68(2):102-107 February 1990
Sesumum portulacastrum L., Syst. Nat., ed. 10. 1058. 1759.
Portulaca portulacastrum L., Sp. Pl 446. 1753.
Sesuvmum revolutifokum Ortega, Hort. Matr., Dec. 19. 1797. Sesumum ortegae Spreng., Nachr. Bot. Gart. Halle 1:36. 1801. Sesummum pedunculatum Pers., Syn. Pl. 2:39. 1806.
Sesuvium sessile Pers., Syn. Pl 2:39. 1806.
Sesuvtum sesstlflorum Domb. ex Rohrb., en Martius Fl Bras. 14, 2:310. 1872.
Hahmus portulacasitrum O. Kuntze, Rev. Gen. 1:263. 1891.
Nombre comun: Ts’a’ay kaan, ts’in kaan, xaw tsikin, xukul, pico real, verdolaga xukul, verdolaga de costa, verdolaga de la playa.
Hierba perenne; tallo rastrero o erecto, muy ramificado, hasta 2 m de diametro, de color verde-morado, liso, glabro. Hojas oblongas a oblongo- lanceoladas, suculentas, 0.7-5 cm de largo y 0.2-0.6 cm de ancho, lisas, el margen entero, el apice agudo-obtuso, sin nervacion aparente. pecioladas. Flor pedicelada, el pedicelo de 0.6-1.5 cm de largo, la simetria radial; perianto pen- talobulado, los tépalos ovado-lanceolados a lanceolados, persistentes, la super- ficie interna de color rosa-morada, ca 0.4-0.8 cm de largo y 0.2-0.3 cm de ancho; estambres numerosos, insertos en el tubo floral, perigineos, libres o subunidos, las anteras oblongo-elipticas, alrededor de 0.1 cm de largo, el filamento de 0.3 cm de largo; ovario 3 o 4 carpelar, trilocular, ovoide-globoso, los ovulos nu- merosos, la placentacion axilar; estilos 3 o 4, ca 0.2 cm de largo, el estigma longitudinal, papiloso. Fruto capsular, conico, verde a moreno claro, 0.9-1.1 cm de largo y 0.5-0.6 cm de ancho; semillas ca 50, lenticular-reniformes, 1.2-1.5 mm de diametro, el arilo membranoso, negras, lisas, lustrosas, el endospermo suave, albuminoso, el embrion anular, los cotiledones oblongos, carnosos.
Sesuvtum portulacastrum se distribuye desde los Estados Unidos hasta Su- damerica a lo largo de las costas de los Oceanos Atlantico y Pacifico. En Mexico a lo largo de ambas costas e islas cercanas y en algunos lugares del interior (zonas secas o salinas). En la Peninsula de Yucatan se distribuye a lo largo de toda la costa, siempre al nivel del mar, en manglares y vegetacion de duna costera. Florece de febrero a noviembre, siendo mas intensa en el verano. La apariencia de esta especie es muy variable, dependiendo del medio donde se la encuentre. Generalmente se presenta mejor desarrollada en las dunas, los individuos en los manglares ocupan zonas extensas pero son pequenos en sus estructuras, v.g., largo de hojas, ancho del tallo. El verano es la epoca del ano donde se presenta el maximo desarrollo de esta especie.
Ejemplares examinados.
Rico-Gray: Aizoaceae de la Peninsula Yucatan, México 105
Yucatan: Telchac Puerto, Calzada, et al. 6603 (XAL); Isla Blanca de Ar- recife Alacranes, Chan 830 (XAL); entrada al puerto de Sisal, Chan 2837 (XAL); Rio Lagartos junto al ojo de agua, Chan y Puch 1544 (XAL); 15 km de Celestun camino a Sisal, Espejel 47(XAL); Isla Pérez en Arrecife Alacranes, Flores y Ucan 9231 (XAL); Isla Larga de Arrecife Alacranes, Flores y Ucan 9249 (XAL); Isla Muertos en Arrecife Alacranes, Narvaez, et al. 755 (XAL); Celestun, Palma y Allkin 281 (XAL); 6 km al W de Sisal, Puch y Espejel 582 (XAL); Rio Lagartos a orilla del ojo de agua, Puch y Chan 808 (XAL); 800 m al S de Celesttin orilla del estero, Rico-Gray 63 (XAL); 4 km al S de Sisal, Rico-Gray 76 (XAL); 1 km al S de Chelem, Rico-Gray 83 (XAL); 1.5 km al E de Dzilam de Bravo, Rico-Gray 92 (XAL); Rio Lagartos, Rico-Gray 108 (XAL); Sisal, Rico-Gray 354 (XAL).
Campeche: entre Villa Madero y Seybaplaya, Calzada, et al. 6782; km 4 carr. Campeche-Champoton, Chan y Burgos 673 (XAL); Isla Centro de Ar- recife Cayo Arcas, Flores y Ucan 9181 (XAL); camino a El Zapote cerca de Champoton, Puch 47; camino a El Zapote cerca de Champoton, Rico-Gray 139 (XAL); Isla Arena, Rico-Gray 225 (XAL); Isla del Centro de Cayo Arcas, Zamudio 100 (CSAT,XAL).
Quintana Roo: lado SE de Isla Mujeres, Chan, et al. 1568 (XAL); lado SE de Isla Contoy, Chan, et al. 1626 (XAL); lado N de Isla Holbox, Chan, et al. 1644 (XAL); parte S Isla Cozumel, Chan, et al. 1763 (XAL); Isla Mujeres, Flores y Ucan 8376 (XAL); Isla Contoy, Flores y Ucan 8845, 8861 (XAL); Isla Holbox en Cabo Catoche, Flores, et al. 9371 (XAL); parte E de Isla Mujeres, Flores, et al. 9271 (XAL); parte W de Isla Contoy, Flores, et al. 9334; islote norte de Cayo Culebras, Flores, et al. 9522(XAL); playa Ojo de Agua en Pto. Morelos, Moreno 621(MEXU); playa de Pta. Brava 4 km al S de Pto. Morelos, Moreno 844 (MEXU); Cayo Centro de Banco Chinchorro, Narvaez y Ucan 891 (XAL); Cayo Norte de Banco Chinchorro, Narvaez y Ucan 906 (XAL); lado W de Isla Mujeres, Puch, et al. 840 (XAL); lado E de Isla Contoy, Puch, et al. 903 (XAL); 4 km al W de Cabo Catoche, Puch, et al. 938 (XAL); Vigia Chico, Rico-Gray 134 (XAL); entre Pto. Juarez y Punta Sam, Rico-Gray y Chan 399 (XAL); parte N de Isla Holbox, Rico-Gray 175 (XAL); Playa Maya al S de Isla Cozumel, Rico-Gray 251 (XAL).
TRIANTHEMA L., Sp. Pl. 1:223. 1753.
Hierbas suculentas, erectas o postradas, ramificadas. Hojas opuestas, las del mismo par de distinto tamano (una mas grande que la otra), pecioladas, la base del peciolo envolvente. Flores pequenas, sésiles, generalmente solitarias en las axilas de las hojas; perianto pentalobulado, concavo, la superficie interior de color rosa muy tenue; estambres 5-10, perigineos, alternos a los sépalos cuando igual en numero; ovario semi-infero, 1-2 léculos, pocos ovulos; estilos 1-2. Fruto capsular, membranoso o coriaceo, con apéndices alados prominentes en
106 PHY POLE GIA volume 68(2):102-107 February 1990
el apice, carnoso, circuncisil longitudinalmente; semillas reniformes, el embrion anular, los cotiledones oblongos.
Se presentan alrededor de 15 especies en los tropicos o en regiones calidas de ambos hemisferios (Nevling 1961). Solo una especie se presenta en Norteamerica
(Correll & Johnston 1970).
Trianthema portulacastrum L., Sp. Pl. 223. 1753.
Trianthema monugyna L., Gen. Pl. 105. 1754.
Trianthema monogynum L., Mant. 69. 1767. Portulacastrum monogy-
num (L.) Medic., Phil. Bot. 1:99. 1789. Trianthema procumbens Mill., Gard. Dict., ed. 8. No. I. 1768. Trianthema fleruosa Schum. & Thorn., Beskr. Guian. Pl. 221. 1828.
Nombre comun. Verdolaga.
Hierbas anuales, rastreras o erectas, formando carpetas hasta de 1 m; tal- los muy ramificados, los jovenes pubescentes. Hojas obovadas o suborbicular- elipticas, suculentas, 0.7-2.8 cm de largo y 0.3-1.3 cm de ancho, lisas, el margen entero, el apice agudo a obtuso, la base cuneada rodeando al tallo, solo la ner- vadura central aparente, ocasionalmente el borde de la hoja de color morado; peciolo de 0.3-0.7 cm de largo. Flores axilares, solitarias, la simetria radial; perianto pentalobulado, los lobulos ovado-lanceolados a lanceolados, la super- ficie interior de color rosado (muy tenue), ca 0.25 cm de largo; estambres 5-10, insertos en el orificio del tubo del perianto, perigineos, el filamento filiforme, ca 0.15 cm de largo, glabro, las anteras ovoideas, ca 0.05 cm de largo e igual de ancho; ovario unicarpelar, turbinado, de 0.01 cm de largo e igual de diametro, glabro, uno o dos loculos, pocos ovulos, placentacion parietal, estilo de 0.01 cm de largo. Fruto capsular, cilindrico, verde a moreno obscuro, 0.4-0.5 cm de largo; semillas 2-5, reniformes, ca 0.2 cm de diametro, la testa rugosa, negra- rojiza, el endospermo albuminoso, el embrion anular, los cotiledones oblongos, carnosos.
Trianthema portulacastrum se distribuye desde el sur de Estados Unidos hasta Sudamerica. En México, en ambas costas y en algunos lugares del inte- rior. Es una planta muy escasa en la peninsula, solo ha sido colectada, entre el nivel del mar y los 20 m, en tres localidades del Estado de Yucatan, en manglares y como planta secundaria en basureros y en zonas arqueologicas. Florece todo el ano.
Ejemplares examinados.
Yucatan: Chichen Itza, Gaumer 743 (MO); Izamal, Lundell & Lundell 8207 (MICH); Sisal, Reco-Gray 350 (XAL).
Rico-Gray: Aizoaceae de la Peninsula Yucatan, México 107
LITERATURA CITADA
Bogle, A.L. 1970. The genera of Molluginaceae and Aizoaceae in the South- eastern United States. J. Arnold Arbor. 51:431-462.
Correll, D.S. & H.B. Correll. 1972. Aquatic and Wetland Plants of South- western United States. (Aizoaceae). Environmental Protection Agency.
U.S. Government Printing Office. Washington, D.C. p. 870-879.
Correll, D.S. & M.C. Johnston. 1970. Aizoaceae. in: Manual of the Vascular Plants of Teras. Texas Research Foundation. Renner, Texas. p. 602-605.
Lot-Helgueras, A. & V. Rico-Gray. 1978. Nota sobre el registro de Sesuvium maritimum ( Walt.) B.S.P. (Aizoaceae), en las costas de México. Biotica 3(1):25-28.
Lundell, C.L. & A.A. Lundell. 1983. The flora of northern Yucatan and the Coba area of Quintana Roo, Mexico: collections and observations in
1938. Wrightia 7(3):97-228.
Nevling, L.I., Jr. 1961. Aizoaceae. in: Flora of Panama. Ann. Missouri Bot.
Gard. 48(1):80-85.
Raven, P.H. & D.L. Axelrod. 1974. Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Gard. 61(3):539-673.
Rico-Gray, V. 1979. Aizoaceae. En: A. Gomez-Pompa & V. Sosa, (eds.), Flora de Veracruz. Fasciculo 9. INIREB, Xalapa, Veracruz, México. 13
PP.
Rzedowski, G.C. 1979. Aizoaceae. En: J. Rzedowski & G.C. Rzedowski, (eds.), Flora Fanerogdmica del Valle de Mézico. CECSA. Mexico. v. 1. p. 162-163.
Sosa, V., J.S. Flores, V. Rico-Gray, R. Lira & J.J. Ortiz. 1985. Lista Floristica y Sinonimia Maya. En: V. Sosa, (ed.), Etnoflora Yucatanense. Fasciculo 1. INIREB, Xalapa, Veracruz, México. 225 p.
Standley, P.C. 1930. Flora of Yucatan. Field Mus. Nat. Hist., Bot. Ser. III(3):157-492.
Standley, P.C. & J.A. Steyermark. 1946. Aizoaceae. En: Flora of Guatemala. Fieldiana, Bot. 24(4):203-207.
Phytologia (February 1990) 68(2):108-112.
BATACEAE DE LA PENINSULA DE YUCATAN, MEXICO
Victor Rico-Gray Centro de Ecologia, UNAM, A.P. 70-275, 04510 México, D.F. MEXICO
RESUMEN
Se presenta la descripcion de la familia Bataceae (=Batidaceae) para la Peninsula de Yucatan, México (Estados de Yucatan, Campeche y Quintana Roo). Se describe taxonomicamente a Batis maritima; se presenta informacion sobre su distribucion y ejemplares examinados.
ABSTRACT
A description of Bataceae (=Batidaceae) from the Yucatan Penin- sula of México (states of Yucatan, Campeche and Quintana Roo) is pre- sented. A taxonomic description of Batis maritima, information about its distribution and a list of specimens examined are presented.
PALABRAS CLAVE (KEY WORDS): Bataceae (=Batidaceae),
Batis maritima, Yucatan, Campeche, Quintana Roo, México.
INTRODUCCION
Los miembros se la familia Bataceae Meiss. (=Batidaceae) se caracterizan por ser arbustos o sufrutices, monoicos o dioicos, hasta de 1.5 m de altura, con tallos postrados o erectos, suculentos; hojas simples, opuestas, decusadas, sésiles, lineares a claviformes, suculentas, glabras, enteras; estipulas caducas, pequenas; inflorescencias masculinas axilares o terminales, conicas 0 espigadas con las flores en las axilas de bracteas; flores con 4 estambres alternisépalos, estos insertos en la base del perianto, los filamentos libres, engrosados o lin- eares y alados; inflorescencias femeninas axilares, con las flores sostenidas por bracteas y agregadas en conos; flores aperiantadas; ovario bicarpelar, sésil, 4-loculado, con un 6vulo anatropo en la base de cada léculo, el pericarpio esponjoso; semillas sin endospermo, la testa membranosa; embrion erecto, los cotiledones largos (Rico-Gray & Nee 1982).
Se situa el origen de la familia en el Cretacico Superior (Maestrictiano) (Muller 1981). En el mundo esta familia esta representada por un solo género con dos especies: Batis maritima L. en las costas tropicales y subtropicales
108
Rico-Gray: Bataceae de la Peninsula Yucatan, México 109
del Nuevo Mundo y B. argillicola van Royen, en la costa sur de Nueva Guinea (Van Royen 1956). Las dos especies son muy similares, B. argullicola difiere principalmente en que es monoica con las flores sostenidas (envueltas) por hojas y no agregadas en conos (espigas cortas); tambien es muy similar en habito a B. maritima y crece, asi mismo, en regiones salinas cercanas al mar.
La situacion filogenetica del género Batis es un tanto incierta, por una parte ha sido asociado con el orden Caryophyllales (Subclase Caryophillidae) por la apariencia y habitos similares al género Sarcobatus (Chenopodiaceae) (Mabry & Turner 1964; McLaughlin 1959). En general, Batzs tiene ciertas car- acteristicas que no estan presentes en ese orden (Caryophyllales), como son: la ausencia de antocianinas y betalaninas, embrion recto en lugar de curvo, ovario tetralocular y un ovulo por loculo. Morfologicamente se puede rela- cionar con los Hammamelidae aunque no esta emparentado. La incertidumbre en la ubicacion filogenética de las Batales, también la comparten otros autores (Kagan & Mabry 1969), quienes indican que fitoquimicamente sus afinidades son muy oscuras como las de Caryophyllaceae y Molluginaceae. Varias lineas de evidencia indican que la familia puede estar relacionada al orden Cappar- ales. Con este comparte la presencia de mirosinasa (Schraudolf, et al. 1971) y también existe evidencia a favor por el numero de cromosomas (Goldblatt
1976). BATIS L., Syst. Nat., ed. 10. 1289, 1380. 1759. Con los mismos caracteres de ia familia. Batis maritima L., Syst. Nat., ed. 10. 1289, 1380. 1759.
Nombre comun: Alambrillo, saladillo.
Arbustos o sufrutices, generalmente muy ramificados, hasta de 60 cm de altura; tallos erectos, los mas viejos postrados, teretes, hasta de ca 6 mm de diametro, los jovenes amarillo-verdosos, verde palidos al secarse, suculentos, glabros. Hojas de 0.5-4 cm de largo por 0.5-1 cm de ancho cuando secas, suculentas, glabras, el apice agudo o diminutamente apiculado, la base pro- longada mas alla del punto de insercion en el tallo para formar un lobulo, este redondeado a agudo o ligeramente trilobado, de ca 1 mm de largo, recur- vado al secar. Inflorescencias masculinas axilares, en forma de espiga, sésiles a subpedunculadas, de 5-10 mm de largo por 3-4 mm de ancho; bracteas en igual numero que las flores, en 4 filas, imbricadas, persistentes, de ca 2 mm de largo y de ancho, obtusas a apiculadas; flores 12-32; caliz en forma de copa, bilobado; corola ausente; estambres 4, insertos en la base del caliz, los fila- mentos de ca 1.5 mm de largo, aplanados cuando secos, alternos y mas largos que los estaminodios triangulares, las anteras de 0.8-1.2 mm de largo, dorsifi- jos. Espigas femeninas con 4-12 flores; sin caliz ni corola, pero sostenidas por pequenas bracteas deciduas, suborbiculares, ca 2 mm de ancho, apiculadas;
110 PHY TOLO EDA volume 68(2):108-112 February 1990
pistilos unidos en la mitad inferior para formar un conjunto, de ca 6 mm de largo por 2.5 mm de grueso, papilado-puberulentos, el apice libre, altusado y dirigido hacia la espiga, los estigmas bilobados, sesiles. Fruto mas o menos ovoide, irregular, con los estigmas persistentes, de 5-15 mm de largo por 5 mm de grueso.
Batis maritima se distribuye en ambas costas del Continente Americano, desde California hasta Peru en el oeste y desde Florida hasta Brasil en el este (Burger 1977; Correll & Correll 1975; Correll & Johnston 1970; Mason 1957; Shreve & Wiggins 1964; Standley 1930). Es comun en la mayoria de las Islas del Caribe. Ha sido introducida en las Islas Hawaii. En Mexico se presenta a lo largo de los dos litorales, generalmente asociada a manglares. En la Peninsula de Yucatan se presenta a lo largo de la costa de los tres estados, asociada a manglares; en general terrenos que son inundados por agua salina durante una epoca del ano y que al secarse dejan suelos con altas concentraciones salinas. El habito de la planta es muy variable dependiendo de las condiciones de salinidad y nivel de agua del suelo; podemos encontrar plantas totalmente postradas y otras erectas. Florece de marzo a septiembre.
Ejemplares examinados.
Yucatan: Progreso, Bruff 1473 (MEXU); 2 km al W de Sisal, Chan y Rico- Gray 1546(XAL); camino El Cuyo-Colonia Yucatan, Narvaez 26 (XAL); carr. Telchac Puerto-Chicxulub Pto., Narvaez y Espejel 644 (XAL); Sta. Clara- Dzilam de Bravo, Palma y Allkin 307 (XAL); orilla del ojo de agua en Rio Lagartos, Puch y Chan 812(XAL); 800 m de Celestun a orilla del estero, Rico- Gray 70 (XAL); 1 km al S de Chelem, Rico-Gray 82 (XAL); 1.5 km al E de Dzilam de Bravo, Rico-Gray 97(XAL); Rio Lagartos, Rico-Gray 105 (XAL); 3 kms al S de El Cuyo, Rico-Gray 111 (XAL).
Campeche: carr. Villahermosa-El Zacatal, Bonilla 46 (ENCB); Punta Are- nas, Chan 416 (XAL); El Remate, Chan y Burgos 662 (XAL); Punta Arena, Chan y Flores 416 (XAL); Pto. Real a Cd. del Carmen, Chan y Flores 509 (XAL); El Zacatal, Chavelas y Zamora ES-4753 (MEXU)); isla del centro en Arrecife Cayo Arcas, Flores y Ucan 9187 (XAL); 6 km antes de la costa en camino Pomuch-Isla Jaina, Gongora 558 (XAL); 1 km antes del puente de Sa- bancuy, Gongora 577(XAL); Cd. del Carmen, Mftranda 7946 (MEXU); camino a El Zapote cerca de Champoton, Puch 46 (XAL).
Quintana Roo: 2 km antes del Mirador Maya en Isla Mujeres, Chan, et al. 1591 (XAL); Isla Contoy, Chan, et al. 1634 (XAL); Holbox, Flores 8497 (XAL); parte E de Isla Mujeres, Flores y Ucan 8403 (XAL); Isla Mujeres, Flores y Ucan 8424, 8811 (XAL); Holbox, Flores y Ucan 8497 (XAL); Cayo Centro en Banco Chinchorro, Flores y Ucan 8965, 8973 (XAL); Isla Mujeres, Flores, et al. 9299 (XAL); Isla Contoy, Flores, et al. 9344 (XAL); Cayo Cule- bras, Flores, et al. 9511 (XAL); Cayo Centro en Banco Chinchorro, Narvaez
Rico-Gray: Bataceae de la Peninsula Yucatan, México 111
809 (XAL); lado E de Isla Contoy, Puch, et al. 912(XAL); Vigia Chico, Rico- Gray 135 (XAL); Holbox, Rico-Gray 203 (XAL); Chiquila, Ucan 430 (XAL); Cayo Norte en el Banco Chinchorro, Ucan 2099 (XAL); atras del Puerto de Abrigo de Isla Mujeres, Ucan y Flores 1040, 1050, 1051 (XAL); Isla Mujeres, Ucan y Flores 1079 (XAL); ruinas de La Mundaca en Isla Mujeres, Ucan y Flores 1239 (XAL).
LITERATURA CITADA
Burger, W. 1977. Bataceae. En: Flora Costarricensis. Fieldiana, Bot. 40:54-55.
Correll, D.S. & H.B. Correll. 1975. Bataceae. in: Aquatic and Wetland Plants of the Southwestern United States. Stanford University Press. p. 868-870.
Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of
Teras. Texas Research Foundation, Renner, TX.
Cronquist, A. 1960. The Evolution and Classification of Flowering Plants. Houghton Mifflin, Co. Boston.
Goldblatt, P. 1976. Chromosome number and its significance in Batis mar- ituma (Bataceae). J. Arnold Arbor. 57:526-530.
Kagan, J. & T.J. Mabry. 1969. Isorhamnetin 3-0-rutinoside flavonoid pig- ment in Batis marituma. Phytochemistry 8:325-326.
Mabry, T.J. & B.L. Turner. 1964. Chemical investigations of the Batidaceae. Taxon 13:197-200.
Mason, H.L. 1957. A Flora of the Marshes of California. Univ. of California Press. p. 878.
McLaughlin, J. 1959. The woods and flora of the Florida Keys: wood anatomy and phylogeny of Batidaceae. Tropical Woods 110:1-15.
Muller, J. 1981. Fossil pollen records of extant angiosperms. The Botanical Rev. 47(1):1-142.
Rico-Gray, V. & M. Nee. 1982. Bataceae. En: A. GOmez-Pompa & V. Sosa, (Eds.), Flora de Veracruz. Fasciculo 21. INIREB, Xalapa, Veracruz, Mexico. 6 p.
112 PaY PTeeoeira volume 68(2):108-112 February 1990
Schraudolf, H., B. Schmidt & F. Weberling. 1971. Das Vorkommen von myrosinase als hin Weis auf die Systematische stellung der Batidaceae.
Experientia 27:1090-1091.
Shreve, F. & I.L. Wiggins. 1964. Batidaceae. in: Vegetation and Flora of the Sonoran Desert. 1:489-490. Stanford University Press, Palo Alto.
Standley, P.C. 1930. Flora of Yucatan. Field Mus. Nat. Hist., Publ. 279, Bot. Ser. 3(3):157-492.
Van Royen, P. 1956. A new Batidaceae, Batis argillicola. Nova Guinea n.s. 7:175-195.
Phytologia (February 1990) 68(2):113-119.
RHIZOPHORACEAE DE LA PENINSULA DE YUCATAN, MEXICO
Victor Rico-Gray Centro de Ecologia, UNAM, A.P. 70-275, 04510 Mexico, D.F. MEXICO
RESUMEN
Se presenta la descripcion de la familia Rhizophoraceae para la Peninsula de Yucatan, México (Estados de Yucatan, Campeche y Quin- tana Roo). Se describe taxonomicamente a Cassipourea gutanensis y a Rhizophora mangle; se presenta informacion sobre su distribucion y ejemplares examinados.
ABSTRACT
A description of Rhizophoraceae from the Yucatan Peninsula of México (states of Yucatan, Campeche and Quintana Roo) is presented. Taxonomic descriptions of Cassipourea guianensts and Rhizophora man- gle, along with information about their distribution and a list of speci- mens examined are presented.
PALABRAS CLAVE(KEY WORDS): Rhizophoraceae, Cassipourea guianensis, Rhizophora mangle, Yucatan, Campeche, Quintana Roo, México.
INTRODUCCION
Los miembros de la familia Rhizophoraceae R. Brown se caracterizan por ser arboles 0 arbustos perennifolios, con hojas por lo comun opuestas, estipulas amplias, interpeciolares y caducas; inflorescencias generalmente axilares con flores perfectas, el caliz persistente, los pétalos libres después de la antesis, alternisépalos, en numero doble o igual a aquellos, los estambres libres, el ovario generalmente infero, algunas veces semi-infero o supero, los ovulos 4- muchos, anatropos; fruto una baya o capsula; germinacion a veces vivipara (Gregory 1958; Vazquez-Yanes 1980).
El origen de esta familia se situa en el Eoceno Superior. Algunos autores consideran que el centro de origen se localiza en el sureste asiatico (Chapman 1975; Steenis 1962) y, otros, proponen al antiguo continente de Gondwana como el lugar de origen de un grupo de géneros de la familia (Raven & Axelrod
113
114 PUY T3Q:- LjQiG kA volume 68(2):113-119 February 1990
1974). Es muy probable que futuras investigaciones indiquen que la segunda idea sea la mas acertada, ya que algunos de los restos fosiles mas antiguos del genero Rhizophora provienen de zonas que antiguamente formaron parte de Gondwana (v.g., India).
En America se presentan 3 géneros de esta familia, los cuales por las difer- encias que presentan en el habitat y forma de dispersion los podemos separar en dos grupos. Por un lado tenemos a Rhizophora, su dispersion es por medio de corrientes marinas y habita zonas costeras y, por otro lado, tenemos a los géneros Anisophylla y Cassipourea cuya dispersion se realiza en tierra y habitan bosques tropicales desde el nivel del mar hasta los 2000 msnm. La presencia de los dos ultimos géneros en America se debe a que este continente formaba parte de Gondwana y al separarse este, las partes conservaron gran cantidad de los taxa presentes, lo que se evidencia con la presencia de estos dos géneros en Sudamérica y Africa. Con respecto al género Rhizophora, se piensa que lleg6 a América a través del antiguo Mar de Tetis (Chapman 1975).
La familia Rhizophoraceae es bastante conocida pues contiene a las especies de mangles mas conspicuas. Esta constituida por 16 géneros y 120 especies, la ultima cifra puede variar dependiendo del autor. Se distribuye en las regiones tropicales y subtropicales del mundo, generalmente entre los 25° N y 25° S de latitud. El género con la mas amplia distribucion es Rhizophora. En Mexico se presentan los géneros Cassipourea y Rhizophora (Standley 1930; Standley & Williams 1963).
CLAVE PARA GENEROS DE RHIZOPHORACEAE DE YUCATAN
Clave para los géneros de la familia Rhizophoraceae presentes en la Peninsula de Yucatan:
1. Plantas con raices subterraneas; hojas subcoriaceas; 15-20 estambres; fruto fedora Senn NA IRE a arn ys) nn hs oh site wale bain Cassipourea
1’ Plantas con raices adventicias aereas y raices “zanco;” hojas coriaceas; estambres 8; fruto una baya de la que pende frecuentemente el hipocotilo et SCR HCRAECIMEIS ot ce hc ec vee tee sees ecens Rhizophora
CASSIPOUREA Aubl., Hist. Pl. Guian. Fr. 1:528. 1775.
Tita Scop., Introd. 219. 1777.
Legnotis Swartz, Prod. 84. 1788.
Richaeza Thouars, Gen. Nov. Madag. 25. 1806.
Weihea Spreng., Syst. 2:594. 1825.
Austrutheria Gardn., Calcutta J. Nat. Hist. 6:344. 1846. Dactylopetalum Benth., J. Linn. Soc., Bot. 3:79. 1858.
Rico-Gray: Rhizophoraceae de Ja Peninsula Yucatan, México 115
Richea O. Kuntze, Rev. Gen. 235. 1891.
Arboles o arbustos mas o menos glabros. Hojas opuestas, subcoriaceas o membranosas, el margen entero a crenado o dentado; estipulas interpeciolares pequenas, caducas. Flores en las axilas de las hojas, perfectas, solitarias a fasciculadas, pedunculadas o subsésiles; caliz coriaceo, 4-7 lobulos triangulares, erectos; corona con 4-7 pétalos blanco-cremosos, alternisépalos, insertados en la base de un disco cupular crenado, glabros o pubescentes, caducos; estambres 8-40, los filamentos iguales o mas largos que el caliz; ovario supero, 2-4 locular, 2 ovulos por léculo. Fruto una capsula carnosa, coriacea; 2-4 semillas ariladas.
Este género consta de 60 especies, la mayoria se encuentran en Centro, Sudamerica y Africa; también se encuentra en Madagascar, India, Sri Lanka y Australia. En México solo existe Cassipourea guianensis (Vazquez- Yanes
1980). Cassipourea guianensis Aubl., Hist. Pl. Guran. Fr. 1:529. 1775.
Legnotis elliptica Sw., Prodr. 84. 1788. Cassipourea elliptica Poir., Encycl., suppl. 2, 131. 1811.
Cassipourea podantha Standl., Field Mus. Nat. Hist., Bot. Ser. 4:241. 1929.
Cassipourea macrodonta Standl., l.c. 242.
Cassipourea belizensis Lundell, Bull. Torrey Bot. Club 66:598. 1939.
Nombre comun: Nanchillo negro (Tabasco), waterwood (Belice).
Arbol o arbusto perennifolio; corteza gris o pardo-grisacea, inmumento amarillo claro. Hojas ovadas a elipticas de 2.5-12 cm de largo y 1.4-4.5 cm de ancho, membranaceas, margen entero o dentado ligeramente, apice agudo, base obtusa; peciolos hasta 0.6 cm de largo; nervadura central bien perceptible en el envés de la hoja; estipulas interpeciolares caducas; yema terminal de 0.2-0.4 cm de largo. Flores axilares, fasciculadas o generalmente geminadas, con 2 6 3 flores, pedicelos articulados de 0.2-0.5 cm de largo, hermafroditas, 0.7-0.9 cm de diametro; caliz pentalobulado, persistente, de 0.3-0.4 cm de longitud, coriaceo, glabro; corola con 5 pétalos caducos, blanco-amarillentos, los margenes con vellosidades, laciniados, de 0.5-1.0 cm de largo y 0.4-0.6 cm de ancho; estambres 15-20, blancos, hasta de 0.2 cm de longitud; ovario trilocular, estilo filamentoso de 0.2-0.4 cm de longitud, persistente. Fruto, una capsula de color moreno al madurar, redondeada a ligeramente eliptica, de 0.5-1.1 cm de longitud y 0.3-0.7 cm de diametro.
Cassipourea guianensis habita en las regiones himedas de Centroamerica y sureste de México. Se encuentra también en las Islas del Caribe y norte de Sudamerica hasta Brasil y Peru. Para México se tienen registros para el limite entre los estados de Chiapas y Tabasco, el sur de Veracruz y la Peninsula
116 Pir FOR QCA volume 68(2):113-119 February 1990
de Yucatan. En la peninsula solo se le encuentra al nivel de! mar, hasta unos kilometros al norte del Rio Hondo en el Estado de Quintana Roo, en areas pantanosas formando el sotobosque de una comunidad de Pinus carzbaea. Florece entre abril y noviembre.
Ejemplares examinados.
Quintana Roo: sabana del Jaguacatal ejido de Caobas, Calzada 10535 (XAL); ejido de Caobas, Flores 10194 (XAL); El Jaguacatal ejido de Caobas, Flores y Puch 9064 (XAL).
RHIZOPHORA L., Gen. Pl., ed. 1. 137. 1737.
Mangle Pluk. er Adans., Fam. Pl. 2:445. 1763. Manguim Rumph. ez Scop., Intr. Hist. Nat. 218. 1777. Asophora Neck., Elem. Bot. 2:361. 1970.
Arboles o arbustos monoicos; tronco y ramas apoyados en raices adventi- cias. Hojas simples, pecioladas, glabras, coriaceas, margen entero; estipulas interpeciolares, sésiles, lanceoladas, foliosas, recubriendo a las hojas jovenes antes de la expansion, luego caducas. Inflorescencias axilares, cimosas; pedun- culos dicotomicamente ramificados; flores 2-20 6 mas por inflorescencia, perfec- tas; sepalos 4, libres, persistentes, coriaceos, glabros; petalos 4, alternisépalos, libres, caducos, coriaceos o membranosos, cubiertos por pelos simples; estam- bres 8 6 12, las anteras subsésiles, multiloculares; ovario semi-infero, cada loculo con 2 ovulos anatropos suspendidos del apice. Fruto una baya coriacea; una semilla con el embrion viviparo.
El género presenta 3 especies en el continente americano, de las cuales solo Rhizophora mangle se encuentra en la Peninsula de Yucatan (Breteler 1969;
1977; Rico-Gray 1981). Rhizophora mangle L., Sp. Pl. 1:443. 1753.
Rhizophora americana Nutt., N. Amer. Sylv. 1. 1842.
Rhizophora mangle var. samoensts Hochr., Cand. 2. 1925. Rhizophora samoensis (Hochr.) Salvoza., Nat. Appl. Sci. Bull. Univ. Philipp. 5:179-237.
Nombre comun: Taab che’, taap che’, xtaab che’, mangle colorado, mangle rojo.
Arbol perennifolio, hasta de 15 m; tronco hasta de 30 cm de diametro (d.a.p.); corteza pardo-rojiza, gruesa, fisurada, inodora, amarga; indumento de color rojo intenso; tronco y frecuentemente las ramas apoyados en numerosas raices aéreas de origen adventicio, simples 0 dicotomicamente ramificadas, con
Rico-Gray: Rhizophoraceae de Ja Peninsula Yucatan, México 117
numerosas lenticelas. Hojas decusadas, simples, pecioladas, elipticas a oblon- gas, de 5.5-11.7 cm de largo y 2.4-5.8 cm de ancho, coriaceas, lisas, gruesas, el haz glabro, verde intenso, brillante, el enves glabro, verde claro o amarillento, con puntuaciones pardas, el margen entero, el apice agudo, la base obtusa; nervaduras poco perceptibles en la superficie de la hoja; estipulas interpecio- lares, caducas, caen al extenderse la hoja, foliosas, sésiles, convolutas, con una doble hilera de escamas glandulares en la base interna produciendo un liquido viscoso, las cicatrices foliares persistiendo por 2 anos; peciolos de 1.6-2.7 cm de largo; yema terminal de hasta 5.5 cm de largo, cubierta por las estipulas, de crecimiento continuo, las yemas laterales pequenas. Inflorescencias sim- ples, cimosas o geminadas, con 2 0 3 flores; pedunculos de 2.2-4 cm de largo; pedicelos de 0.4-1.7 cm de largo; flores hermafroditas de 2-2.5 cm de diametro; sépalos 4, persistentes, amarillos, de 9-19 mm de largo por 4-5 mm de ancho, coriaceos, glabros, gruesos; petalos 4, de ca 1 cm de largo por ca 2 mm de ancho, alternisépalos, caducos, blancos o amarillentos, lanceolados, involutos, el margen densamente velloso; estambres 8, subsésiles, multiloculados, de ca 6 mm de largo; estilo bifido de ca 5 mm de largo. Fruto una baya de color pardo, coriacea, dura, piriforme, farinosa, de 1.5-3.5 cm de largo por 0.8-1.9 cm de ancho en la base, caliz persistente; semilla una, rara vez dos, el embrion sin latencia, creciendo en el fruto hasta perforarlo por el apice y emergiendo al ex- terior, el hipocotilo creciendo unido al fruto, desprendiéndose cuando alcanza de 10-31 cm de largo, el endospermo transformandose en un organo placen- tario que permite el intercambio entre el embrion en desarrollo y la planta, los cotiledones fusionados formando un tubo verde recubriendo a la plumula hasta el desprendimiento del embrion, los cotiledones permaneciendo unidos al fruto; el hipocotilo alcanza al desprenderse hasta 31 cm de largo, pardo en la parte inferior y verde en la superior, con numerosas lenticelas, plumula de hasta | cm.
Rhizophora mangle habita esteros, bahias, desembocadura de rios, lagu- nas costeras de las costas de América, en forma continua. En el Oceano Pacifico, desde el sur de Sonora y Baja California hasta Ecuador, incluyendo el Archipiélago de las Galapagos, se le encuentra también en algunas islas de los archipiélagos de Polinesia y Melanesia y ha sido introducido a Hawaii y, en al Oceano Atlantico, se le encuentra en forma discontinua desde la Florida a Brasil, en las Islas del Caribe y en la costa oeste de Africa (Breteler 1969, 1977; Gill & Tomlinson 1969; Graham 1964; Rico-Gray 1981). En la Peninsula de Yucatan se encuentra presente a lo largo de la costa de los tres estados que la componen, ocupando extensas zonas en los manglares y petenes de los estados de Campeche y Quintana Roo. Florece todo el ano, siendo mas intensa la floracion entre abril y julio.
Ejemplares examinados.
Yucatan: Laguna Rosada 1 km antes de llegar a Telchac, Calzada, et al.
118 PHY TOLOGIA volume 68(2):113-119 February 1990
6613 (XAL); Isla Pérez de Arrecife Alacranes, Flores y Ucan 9235 (XAL); puente en el camino Rio Lagartos-Las Coloradas, Leal y Rico-Gray 131 (XAL); camino Sisal-Celestun, Narvaez y Puch 808 (XAL); Rio Lagartos, Ordonez 361 (XAL); 7 km de Celestun hacia Kinchil, Rzco-Gray 62(XAL); 5 km de Celestun a Kinchil, Rico-Gray 71 (XAL); 4 km al S de Sisal, Rico-Gray 77 (XAL); 1.5 km al S de Chelem, Rico-Gray 88 (XAL); 2.5 km al E de Dzilam de Bravo, Rico-Gray 98 (XAL); 1 km al S de San Felipe, Rico-Gray 109 (XAL); cerca de Telchac Pto. en carr. a Motul, Rico-Gray 368 (XAL).
Campeche: zona de los petenes, mpio. de Calkini, Chan 272 (XAL); Isla Punta Arena, Chan 1934 (XAL); Isla Punta Arena, Chan y Flores 426 (XAL); orilla del Rio Zapote, Chan y Yam 1535 (XAL); camino El Remate-Punta Arena, Narvaez y Rico-Gray 247 (XAL); camino a El Zapote, Rico-Gray 140 (XAL); Playa Azul entre Champoton y Sabancuy, Rico-Gray 146 (XAL); camino E] Remate-Isla Arena, Rico-Gray 217 (XAL); orilla del Rio Zapote, Yam 224 (XAL).
Quintana Roo: Boulevard Bahia en Chetumal, Calzada, et al. 7035 (XAL); 4 km de Xcalak en carr. a Majahual, Calzada, et al. 7154 (XAL); Laguna Guerrero, Chan y Burgos 1436 (XAL); lado SE de Isla Mujeres, Chan, et al. 1566, 1619 (XAL); km 38 camino Tulum-Pta. Allen, Duran y Gutierrez 48 (CIQRO,XAL); Juan Sarabia al margen del Rio Hondo, Flores y Ucan 8237 (XAL); a 5 km de Carrillo Puerto, Flores y Ucan 8333 (XAL); Laguna de Chunyaxche, Flores y Ucan 8350 (XAL); orilla de puerto de abrigo en Isla Mujeres, Flores y Ucan 8419 (XAL); Isla Holbox, Flores y Ucan 8476 (XAL); Isla Mujeres, Flores y Ucan 8804 (XAL); Cayo Centro en Banco Chinchorro, Flores y Ucan 8970 (XAL); Islote Cayo Centro en Banco Chinchorro, Flores, et al. 8958 (XAL); Isla Contoy, Flores, et al. 9270, 9327 (XAL); Isla Holbox, Flores, et al. 9366 (XAL); Cayo Culebras islote N, Flores, et al. 9502 (XAL); Cayo Centro en Banco Chinchorro, Narvaez y Ucan 880 (XAL); Cayo Norte de Banco Chinchorro, Narvaez y Ucan 904 (XAL); lado E de Isla Contoy, Puch, et al. 896 (XAL); 4 km al W de Cabo Catoche, Puch, et al. 934 (XAL); entre Pto. Juarez y Cancun, Rico-Gray 119 (XAL); cerca de zona arqueologica de Tulum, Rico-Gray 129 (XAL); Vigia Chico, Rico-Gray 132 (XAL); NE de Isla Holbox, Rico-Gray 207(XAL); Banco Chinchorro, Ucan 2089 (XAL); Isla Mujeres, Ucan y Flores 1235 (XAL); Cayo Centro del Banco Chinchorro, Ucan y Flores 1320 (XAL); Cayo Centro en Banco Chinchorro, Ucan y Narvaez 2058 (XAL); camino Xcalak a Majahual, Ucan, et al. 640(XAL); orilla Rio Hondo, Ucan, et al. 913 (XAL).
LITERATURA CITADA
Breteler, F.J. 1969. The Atlantic species of Rhizophora. Acta Bot. Neerl. 18(3):434-441.
Rico-Gray: Rhizophoraceae de la Peninsula Yucatan, Mexico 119
Breteler, F.J. 1977. America’s Pacific species of Rhizophora. Acta Bot.
Neerl. 26(3):225-230.
Chapman, V.J. 1975. Mangrove biogeography. zn: G. Walsh, et al., (eds.) Proc. Int. Symp. Biol. Managm. of Mangroves Hawaii [:3-22.
Gill, A.M. & P.B. Tomlinson. 1969. Studies on the growth of red mangrove (Rhizophora mangle L.). I. Habit and general morphology. Biotropica 1(1):1-9.
Graham, S.A. 1964. The genera of Rhizophoraceae and Combretaceae in the southeastern United States. J. Arnold Arbor. 45:285-301.
Gregory, D.P. 1958. Rhizophoraceae. in: Flora of Panama. Ann. Missouri Bot. Gard. 45:136-142.
Raven, P.H. & A.I. Axelrod. 1974. Angiosperm biogeography and past con- tinental movements. Ann. Missouri Bot. Gard. 61(3):539-673.
Rico-Gray, V. 1981. Rhizophora harrisoni (Rhizophoraceae), un nuevo reg- istro para las costas de México. Bol. Soc. Bot. Mexico 41:163-165.
Standley, P.C. 1930. Flora of Yucatan. [Field Mus. Nat. Hist., Bot. Ser. 3(3):157-492.
Standley, P.C. & L.O. Williams. 1963. Rhizophoraceae. an: Flora of Guatemala. Fieldiana, Bot. 24(7):263-268.
Steenis, C.G.G.J. van. 1962. The distribution of mangrove plant genera and its significance for paleogeography. K. Ned. Akad. Wet. Proc. Ser. ©, 65:164-169.
Vazquez- Yanes, C. 1980. Rhizophoraceae. En: A. Gomez-Pompa & V. Sosa, (eds.) Flora de Veracruz. Fasciculo 12. INIREB, Xalapa, Veracruz, Mexico. 8 p.
Phytologia (February 1990) 68(2):120-121.
A NEW SPECIES OF AIJCONIA (MELASTOMATACEAE) FROM DOMINICA, LESSER ANTILLES
Alice C. Nicolson 3435 S. 8th St., Arlington, Virginia 22204 U.S.A.
ABSTRACT Miconia mornicola is described from Dominica.
KEY WORDS: Melastomataceae, Muiconia, Lesser Antilles, sys- tematics.
During preparation of Melastomataceae for D.H. Nicolson’s Flora of Do- minica, Part 2, Dicotyledones (in press), a new taxon was detected. Publica- tion was delayed for lack of flowering material and the inability to determine whether the species was dioecious, as expected, or if the flowers were bisexual as they are in the most recent collections.
Miconia mornicola A.C. Nicolson, sp. nov.
A Miconza globuliflora (L. Richard) Cogniaux foliis obtuse cau- diculatis, floribus bisexualibus et seminibus majoribus, laevibus differt. HOLOTYPUS: DOMINICA: Morne Diablotins, “elfin wood- land on ridge dominated by Clusza, alt. 4,500 ft, shrub 2.5 m. Calyx (hypanthium] purple. Petals white, reflexed, soon falling. Stamens splayed; filaments white, anthers cream; dark ring around white ovary. Leaf mid-green, glossy.” 2 Dec 1988, A. Lack, M. Purkins & A. Taylor | (BM); Isotype: US-3145481.
Dense tree to 3 m; young leaves and branches sparsely rusty-scurfy; leaves 5 veined, glabrous, coriaceous, gland dotted beneath, margins dentate with incurved teeth, broadly elliptic, abruptly bluntly caudate, base obtuse-acute, 3.5-5 x 2.5-3 cm; petiole 1-1.5 cm; inflorescence pyramidal-paniculate, to 7 cm; flowers pentamerous, pedicels | mm; hypanthium 1.4 mm, rosy purple, darker within; calyx white, ca 1 mm, tube 0.4 mm, sepals triangular, 0.6 mm, persistent; petals white, round, strongly reflexed, fugacious, 1.7 x 2 mm; stamens 10, 3.5 mm long, broadly spreading at anthesis, filaments 2 mm long, anthers to 1.5 mm, cream, connective with a blunt, basal, bilobed, dorsal tooth to 0.5 mm long and a lobed ventral appendage to 0.2 mm; style exserted before
120
Nicolson: New species of Miconia from Dominica 121
petals open, to 4 mm long, glabrous, stigma scarcely expanded, ovary white, trilocular, 1/3 inferior; fruits globular, bluish, 3 mm; seeds 15-20, smooth, 1.1 x 0.6 mm.
Known only in dwarf forests of Dominican mountain tops: Morne Anglais (Nicolson 4106), Morne Diablotins (Chambers 2643; Lack, et al. 2), Morne Trois Pitons (Ernst 1116A, 2041; Hodge 1420). Flowering early December, fruiting late January (Chambers, Nicolson).
ACKNOWLEDGMENTS
I am grateful to Caroline Whitefoord (BM) who encouraged the collectors, whom I also thank, to search for flowering material. The collectors were Dr. Andrew Lack (Ecology lecturer, Oxford Polytechnic, Oxford, U.K.) and two recent graduates (University of East Anglia, Norwich, U.K.), Melanie Purkins and Andrew Taylor.
Phytologia (February 1990) 68(2):122-12+4.
TWO NEW SPECIES OF ARCHIBACCHARIS (ASTERACEAE: ASTEREAE) FROM OAXACA, MEXICO
Guy L. Nesom Department of Botany, University of Texas, Austin Texas 78713 U.S.A.
ABSTRACT
Two new species of Archibaccharis sect. Archtbaccharis from south central Oaxaca, Méxicoare described: A. macdonaldiiand A. nepho- cephala.
KEY WORDS: Archibaccharis, Asteraceae, Astereae, México.
Recent collections from south central Oaxaca by Dr. Andrew McDonald include a number of previously undescribed species, among them two species of Archtbaccharis. Both are erect shrubs and members of sect. Archibaccharis, but they are only distantly related to each other within that section. Both are known only from the type collection.
Archibaccharis macdonaldii Nesom, sp. nov. TYPE: MEXICO. Oaxaca: Cerro Quiexobra and vicinity, 35 km ESE of Miahuatlan, 5 km NE of Santo Domingo Ozolotepec, timberline vegetation in open glades along ridges and in mountain saddles, dominated below by pine forest, common on dry, ridge tops, SE exposure, 3650-3800 m, 10 Dec 1989, A. AfcDonald 2932 (HOLOTYPE: TEX; isotypes: F,MEXU,NY).
A. hieraciordes (S.F. Blake) S.F. Blake similis sed foliis multo minoribus paucidentatis, capitulis minoribus, et corollis pistillatis ligulis brevibus differt.
Woody, erect shrubs. Stems 1-2 m tall, the upper highly branched and dark purple-brown. Stems, leaves, and phyllaries moderately to densely in- vested with short stipitate resin glands, the stems also sparsely villous. Leaves densely arranged, obovate, basally attenuate to a subpetiolar base, not clasp- ing, 6-16 mm long, 3-7 mm wide, with 1-3 pairs of coarse teeth, thick tex- tured with the upper surface somewhat shiny, glandular but otherwise glabrous or with a few, scattered hairs. Heads campanulate-turbinate, 3-4 mm wide, on wiry pedicels 7-15 mm long, in distinctly flat topped corymbs; phyllaries oblong-lanceolate, purple tipped, graduated, the inner 4.0-4.5 mm long, with
122
Nesom: New species of Archibaccharis from Oaxaca, México 123
hyaline, lacerate-ciliate margins; receptacles deeply alveolate. Pistillate heads: pistillate flowers 28-29, with ligules 0.2-0.3 mm long; hermaphroditic flowers 4, with sterile ovaries; achenes 1.2-1.4 mm long, flat, with 2 thick, lateral ribs, eg- landular, sparsely strigose; pappus bristles 40-45, somewhat uneven in length, the longest 4-5 mm long. Staminate heads not seen.
Archibaccharis macdonald is closely related to A. hteracio1des and A. au- riculata (Hemsley) Nesom (see Nesom 1988) but most similar to the former in its non-clasping, sub-petiolar leaves. With both of the other taxa, the new species shares an erect habit, stipitate glandular vestiture, and relatively large, thin pedicellate heads in an open, distinctly flat topped, corymbose capitulescence. In contrast to both relatives, A. macdonaldw has a woodier habit, thicker, much smaller and coarsely toothed leaves, and smaller heads with ligulate pistillate corollas, as specified in the following couplet.
1. Leaves 4-18 cm long, 20-45 cm wide, with 5-13 shallow, apiculate teeth per side; inner phyllaries 5.0-6.5 mm long; pistillate corollas Bupalalies 5%. dsutleiwndens die wibamedting thepelann din A. hieracioides
1’ Leaves 0.6-1.5 cm long, 3-7 mm wide, with 1-3 coarse teeth per side; inner phyllaries 4.0-4.5 mm long; pistillate corollas with ligules 0.2-0.3 mm Tire, Adamo gadirieu tase be eho hale aere A. macdonald
The only other species of the genus with such extreme woodiness and small, thick leaves is the isolated Archibaccharis peninsularis S.F. Blake from Baja California Sur. Because both A. hieracioides and A. auriculata occur at lower elevations, are more herbaceous, and have much larger and thinner leaves than A. macdonaldi, and because they are similar in these respects to the rest of the genus, I hypothesize that the morphology of the new species is evolutionarily derived with respect to its closest relatives. It is not clear, however, whether it is the sister taxon of A. hieractoides or of both A. hteractordes and A. auriculata.
Archibaccharis nephocephala Nesom, sp. nov. TYPE: MEXICO. Oax- aca: Dirt road between La Cienegilla and San Gregorio Ozolotepec, ca 5 km N of La Cienegilla, roadside weed, pine forest or cloud forest dom- inated by Clethra, Pinus and Quercus, ca 2500-3000 m, 12 Dec 1989, A. McDonald 2971 (HOLOTYPE: TEX; Isotypes: F,MEXU,NY).
A. serratifoliae (Kunth) S.F. Blake similis sed foliis petiolis lon- gioribus laminis longioribus apicibus acuminatioribus et capitulis paucioribus confertim dispositis differt.
Erect shrubs. Stems 2.5 m tall, slightly zig zag in the capitulescence, eglandular, pilose-villous, the hairs with thick and viscid bases. Leaves ovate
124 PUPAE. OOF 1G TA volume 68(2):122-124 February 1990
with long acuminate apices and serrulate-apiculate margins, the blades thin, 9-12 cm long, 3-4 cm wide, basally attenuate to narrow petioles 15-25 mm long, the lower surfaces moderately villous with thin based, crinkly, whitish vitreous hairs, the upper surfaces moderately hispid-pilose with stiffer hairs with thick and orangish bases. Heads cuneate, 3 mm wide, in dense, subcorymboid to rounded panicles; phyllaries lanceolate, graduated, the longest 1.5-2.0 mm long, with thinly pilose-ciliate upper margins, otherwise glabrous or with a few, scattered hairs. Pistillate heads: pistillate flowers 19-20, with ligules 0.4- 0.5 mm long; hermaphroditic flowers 1, with sterile ovaries; achenes 1.0-1.2 mm long, flat, with 2(-4) thick, lateral ribs, eglandular, sparsely strigose to glabrous; pappus bristles 14-16, 1.6-2.0 mm long. Staminate heads: staminate flowers 20-21, the corollas 2.0-2.2 mm long, with abortive ovaries.
The epithet refers to the cumulus shaped capitulescences as well as the cloud forest habitat of the new species.
Archibaccharts nephocephala is most similar, and perhaps most closely re- lated, to the much more widespread A. serrattfolia (Jackson 1975) in its erect habit, eglandular but densely pilose to villous vestiture and ovate-acuminate leaves. The new species differs strongly in its much longer acuminate leaves with longer petioles and in its much smaller and more densely arranged heads with smaller corollas. Technically, they can be separated by the following couplet.
1. Petioles 1-10(-15) mm long; phyllaries 4-5 mm long; staminate corollas
So -Oae tn tone ee eee PPO ARR SP. ERLE, A. serrattifolta
1’ Petioles 15-25 mm long; phyllaries 1.5-2.0 mm long; staminate corollas
ala? shri. Sa Rees OS kk LIE La EG 4. nephocephala ACKNOWLEDGMENTS
I thank Drs. Billie Turner and Andrew McDonald for their review and comments on the manuscript.
LITERATURE CITED
Jackson, J.D. 1975. A revision of the genus Archibaccharis Heering (Com- positae - Astereae). Phytologia 32:81-194.
Nesom, G.L. 1988. Studies in Mexican Archibaccharis (Compositae: Aster- eae). Phytologia 65:122-128.
Phytologia (February 1990) 68(2):125-133.
SYNOPSIS OF OLDENLANDIA (RUBIACEAE) IN THE UNITED STATES
Edward E. Terrell Department of Botany, University of Maryland, College Park, Maryland 20742 U.S.A.
ABSTRACT
A synoptic taxonomic treatment for Oldenlandia (Rubiaceae) is pro- vided for five species occurring in the United States, including data on generic differences, keys to species, synonyms, brief descriptions, and habitats and distributions. Two of the species, O. boscit and O. unt- flora, are native, and three species, O. corymbosa, O. salzmanni and O. callitrichoides, are adventive.
KEY WORDS: Oldenlandia, Rubiaceae, systematics, United States.
This synopsis provides a generic description of Oldenlandia, data on generic differences, keys to species, synonyms, brief descriptions, and habitats and distributions. These data are intended to provide a body of useful taxonomic information for this genus pending completion of a more detailed taxonomic treatment. Five species of Oldenlandia are included, of which two are native to the United States and three are adventive.
Oldenlandia (tribe Hedyotideae; Rubiaceae) is a worldwide genus of warm, subtropical and tropical regions. Verdcourt (1976: 269) estimated the genus to have nearer 100 species, instead of the 300 sometimes estimated (e.g., Willis 1973). Most of the species are native to Africa, with smaller numbers in Asia, Australia and the Americas. Bremekamp (1952) in his extensive monograph of African species of Oldenlandia treated 61 species arranged in 16 subgenera and also dealt with 19 other genera more or less closely related to Oldenlandia. Authors of recent African floras recognized 12 Oldenlandia species in West Tropical Africa (Hepper & Keay 1963), 7 species in Gabon (Hallé 1966) and 37 species in Tropical East Africa (Verdcourt 1976). These authors did not recognize the occurrence of Hedyotis species in their regions, and Hallé (1966) commented that Hedyotis is an Asian genus.
Since 1753, the closely related Linnaean genera Hedyotts, Houstonia and Oldenlandia, have been subjected to the shifting opinions of many system- atic botanists; some of this literature was reviewed by Fosberg (1943) and Bremekamp (1952). Verdcourt (1976: 269), while treating Oldenlandia, cited recent literature on these genera, including papers by Fosberg (e.g. 1943) and
125
126 Pin Y T1010 Gil A volume 68(2):125-133 February 1990
Lewis (1961) that favored the union of these genera under //edyotis. Later how- ever, Lewis (1964, 1965, 1966b) recognized Oldenlandia as a distinct genus, while providing chromosome and pollen data. Verdcourt (1976: 269) com- mented that “If all the genera closely related to Hedyotts are sunk into it, then it forms an unwieldly unit covering a very wide range of structure and habit.” Terrell (1975) expressed a similar viewpoint while pointing out the strong dif- ferences in the type species of the three genera; he did not, however, provide details concerning the total variation. Data concerning seed types, chromo- some numbers and pollen morphology were furnished for 39 North American species of Houstonza (Terrell, et al. 1986). Since 1986 I have concluded that putting most of the North American species in Houstonia is as wrong as placing them all in Hedyotzs. At present, I recognize for North America, approximately 20 species in Houstonza (a genus restricted to North America), 20 species in Hedyotts (with most species in Asia) and 10 species in Oldenlandia (exclud- ing West Indies species). Further data regarding all three genera are being summarized in a subsequent paper.
The bona fide species of Oldenlandia have the following principal charac- teristics: small annual or perennial herbs; corollas small, usually less than 8 mm long, rotate or more or less funnelform or salverform; homostylous or het- erostylous; capsules more or less subglobose, 4/5 to fully inferior; seeds minute (typically 0.2-0.4 mm long), numerous (usually 50-150) per capsule, trigonous or conic in shape, hilum punctiform at apex of an angle, testa minutely retic- ulate, areole walls straight or sinuous, often low and indistinct; chromosome number z=9. The seed characters and chromosome number especially dis- tinguish Oldenlandia from Hedyotis and Houstonia. (The chromosome num- ber z=9 occurs otherwise only in Hedyotts nigricans, which has much larger, compressed-ellipsoid seeds and other morphological differences indicating that it represents a separate phyletic line). Two of the species included here do not have these strictly oldenlandioid seed characters and chromosome num- ber, but they have generally been included in Oldenlandia in the past and one of them (O. salzmanniz) is tentatively retained there pending further study. Both clearly are different from Houstonia and Oldenlandia, so cannot be placed in those genera. The second species, O. callitrichoides Griseb., is being de- scribed by Terrell and W.H. Lewis as a new genus and is included here only to facilitate its identification.
The following generic description is based largely on the species included here (except O. callitrichoides). Also taken into account were the descriptions by Bremekamp (1952) in his monograph of African Oldenlandia and Standley (1918) in his treatment of the genus for the North American Flora.
GENERIC DESCRIPTION
Oldenlandia L., Sp. Pl. 119. 1753. LECTOTYPE SPECIES: O. corymbosa L., selected by Hitchcock & Green (1929).
Terrell: Synopsis of Oldenlandia in United States 127
Small annual or perennial herbs. Stems slender, erect, decumbent or creep- ing. Leaves opposite, sessile or petiolate. Stipules interpetiolar, small. Flowers small, in axillary or terminal glomerules or in cymes or solitary, homostylous (isostylous) or heterostylous (distylous), sessile or pedicellate. Calyx lobes (3-) 4(-5): calyx cup (hypanthium) cup shaped or hemispheric. Corollas 1-5(-8.5) mm long, rotate, funnelform or salverform, white, lavender, pink or purple, glabrous externally; tube shorter or longer than the (3-)4(-5) lobes. Stamens (3-)4(-5), anthers dorsifixed, sessile or on short filaments; filaments inserted on corolla tube at or below corolla sinuses. Stigmas 2 branched; styles filiform or thickened. Capsules small, bilocular, more or less subglobose, fused with calyx cup (hypanthium), 4/5 to fully inferior, loculicidally dehiscent or later also septicidally dehiscent. Seeds usually numerous (50-150) per capsule or in certain species ca 4-30 per capsule, black, brown or tan, minute to small, 0.1-0.4(-0.65 mm) long, usually trigonous or somewhat conic, or in certain species subglobose or oblongoid; hilum punctiform at apex of an angle or at apex of rounded ventral face; testa minutely reticulate; areole walls straight or sinuous, sometimes low and indistinct. Endosperm fleshy: embryo clavate (Standley 1918) or cylindrical. Basic chromosome number in many species r=9, in one species included here n=15.
KEY TO SPECIES
A. Flowers and capsules sessile or on pedicels to ca 3 mm long, | to several in terminal or axillary glomerules.
B. Annual; leaves ovate or elliptic, 3-11 mm wide; calyces and capsules hirsute to glabrous; stipules with conspicuous narrow, marginal, often branched and ciliate, teeth to ca 5 mm long ..2. O. uniflora
B’ Perennial with woody tap root; leaves linear to narrowly elliptic, 1- 3(-5) mm wide; calyces and capsules minutely papillose or verrucose to glabrous; stipules with narrow marginal teeth to ca 2 mm MOET le earn eV Gare ee a's Cn ie ik capac eae ee eee 3. O. bosci
A’ Flowers and capsules on pedicels more than 3 mm long, not in glomerules.
C. Annual, erect, spreading, or decumbent; corollas inconspicuous, more or less obscured by calyx lobes ............ 1. O. corymbosa
C’ Perennial, creeping; corollas small but not obscured by calyx lobes.
D. Leaves broadly elliptic or ovate. 1.5-5.2 mm long, 0.7-2.5 mm wide, sessile or subsessile; corollas pink, purple or white, 2.5-5 mim long; capsules subglobose .............. 4. O. salzmanniu
D’ Leaves suborbicular, subrhombic or broadly ovate, 0.5-4 mm long and wide, petioles 0.5-3 mm long; corollas white, 1.5-3.5
128 PPO bere: 1s volume 68(2):125-133 February 1990
mm long; capsules narrowly turbinate or foci Gries 21. each et EI, Set 5. O. callitrichoides
SYSTEMATIC TREATMENT
1. Oldenlandia corymbosa L., Sp. Pl. 119. 1753. TYPE: To be discussed in a separate paper. Hedyotis corymbosa (L.) Lam., Tabl. Encycl. 1:272. 1792.
Small annual herb. Stems slender, erect, spreading, decumbent or pros- trate, to 2.4(-4) dm tall, usually branched, glabrous or puberulent. Leaves sessile or short petiolate, narrowly elliptic, elliptic, narrowly oblong, or linear, 5-40 mm long, 1-9 mm wide, glabrous or leaf bases ciliate. Stipules to 2 mm long, with 0-few marginal filiform teeth or colleters to 3 mm long. Flowers homostylous, cymose from the axils on slender peduncles to ca 15 mm long, (2-)3(-5) flowers on shorter pedicels from each peduncle, often flowering at most axils. Calyx lobes lanceolate, 0.5-1.3 mm long. Corollas rotate or short funnelform, white or occasionally faint lavender or pink, 1-2 mm long, partly obscured by calyx lobes, corolla tube and lobes about equally long, throat with white hairs. Anthers 0.2-0.3 mm long, attached to lower part of tube. Stigma and style less than 0.5 mm long, style thickened. Mature capsules ]-2.2 mm long, 1.3-2.8 mm wide, subglobose or slightly wider than long, 4/5 to fully inferior, glabrous, truncate or retuse. Seeds numerous per capsule, 0.2-0.4 mm long, trigonous. Flowering all year in tropical climates. Chromosome number: n=9, 18, 27; 2n=18, 36, 54 (Lewis 1959, 1962, 1964, 1966b; and others).
Habitats: Disturbed places, lawns, roadsides. Distribution: Pantropic weed. United States: Mainly in Atlantic and Gulf Coastal Plains and Mis- sissippi Embayment; South Carolina, southern Georgia, throughout Florida, southern parts of Alabama, Mississippi and Louisiana, and rare in eastern Texas. Becoming more common in continental U.S. Hawaii: Oahu, Hawaii, Maui. Also, West Indies and México: Nayarit, Tabasco, Chiapas; probably more common in México than indicated by available records. Standley (1918) did not list any records for México and United States, listing only West Indies, Central America and South America in the Western Hemisphere. Our plants are the diploid race of var. corymbosa (Lewis 1964).
2. Oldenlandia uniflora L., Sp. Pl. 119. 1753. TYPE: To be discussed in a separate paper. Hedyotis unzflora(L.) Lam., Tabl. Encycl. 1:272. 1792. Edrastima uniflora Raf., Actes Soc. Linn. Bordeaux 6:269. 1834.
Hedyotis auricularia Walter, Fl. Carol. 85. 1788. (non H. aurtcularia L.@hn. PL. 101, 1753),
Oldenlandia glomerata Michx., Fl. Bor.-Amer. 1:83. 1803. Hedyotis glomerata (Michx.) Elliott, Sketch Bot. S. Carolina 1:188. 1816.
Terrell: Synopsis of Oldenlandia in United States 129
Stelmotis glomerata Raf., New Fl. 4:101. 1838. Stelmanis glomerata Raf., Autsk. Bot. 13. 1840.
Hedyotis virginica Spreng., Pl. Min. Cogn. Pug. 2:34. 1815.
Hedyotis fasciculata Bertol., Mem. Reale Accad. Sci. Inst. Bologna 2:306. 1850. Oldenlandia fasciculata (Bertol.) Small, Fl. S.E. U.S. 1106. 1903. H. unzflora var. fasciculata (Bertol.) W.H. Lewis, Amer. J. Bot. 49:865. 1962.
Oldenlandia I:ttoralis C. Mohr, Bull. Torrey Bot. Club 24:27. 1897.
Annual herb. Stems slender, erect, spreading or procumbent, to 4.5(-7.7) dm tall (sometimes flowering when plant very small), branched, densely white hirsute or pubescent to glabrous. Leaves sessile or short petiolate, ovate or elliptic, to 28 mm long, 3-11 mm wide, hirsutulous or pubescent to glabrous above or pubescent only on midrib and nerves, glabrous or pubescent on nerves beneath. Stipules to 2 mm long, with 1 to few, sometimes branched and ciliate linear or narrowly lanceolate marginal teeth, to ca 5 mm long. Flowers homostylous, sessile or on pedicels to ca 3 mm long, | to several in axillary and terminal glomerules. Calyx lobes ovate or ovate-lanceolate, 0.8-3 mm long, pubescent to glabrous. Corollas rotate, white or pale blue, inconspicuous, 0.7- 1.3 mm long, usually shorter than calyx lobes. Anthers 0.1-0.2 mm long, attached to corolla tube at sinuses of lobes. Style and stigma 0.2-0.6 mm long; stigmas included in tube; style thickened. Mature capsules 1-2.5 mm long, 1-3 mm wide, subglobose or slightly wider than long, fully or 7/8 inferior, densely hirsute varying to glabrous. Seeds numerous per capsule, 0.2-0.3 mm long, trigonous. Flowering in United States in spring (Florida), summer, fall. Chromosome number: n=18, 36; 2n=36 (Lewis 1962).
Habitats: Wet or moist places, lake shores, swamps, stream banks and sand bars, pine and deciduous woods (often in openings), savannahs, fields, roadsides and gravel pits. Distribution: United States: Mainly in Atlantic and Gulf Coastal Plains and Mississippi Embayment; New York (Long Island), New Jersey, Delaware, Maryland (southeast and Prince Georges Co.), District of Columbia; eastern parts and rare in piedmont of Virginia, North Carolina, South Carolina (incl. Pickens Co.) and Georgia; throughout Florida, Alabama, Mississippi, Louisiana, eastern Oklahoma, eastern Texas, southern Arkansas, southeastern Missouri, western Kentucky and western Tennessee. Also occurs in the West Indies in Cuba, Puerto Rico and Jamaica.
There is some variation in leaf shape and plant vestiture, but these variants, to which names (see synonyms) have been given, definitely vary as continua and I have had no hesitation in sinking these into one species.
3. Oldenlandia bosciz (DC.) Chapm., Fl. Southern U.S. 181. 1860. Hedyotis boscit DC., Prodr. 4:420. 1830. TYPE: UNITED STATES. Carolina: 1798-1800, Bosc s.n. (HOLOTYPE: G-DC [not seen]; Microfiche: US!).
130 PRY POD@WEGrs volume 68(2):125-133 February 1990
Small perennial herb with woody taproot. Stems slender, spreading, de- cumbent or prostrate, to 3 dm tall, or forming mats to ca 4 dm wide, much branched, glabrous to minutely papillose or puberulent. Leaves sessile, linear to narrowly elliptic, to 30 mm long, 1-3(-5) mm wide, glabrous to minutely papillose or puberulent above, glabrous or scabrous on midrib beneath. Stip- ules to 2 mm long, with 1 to several narrow marginal teeth to 2 mm long. Flowers homostylous, sessile or subsessile, 1 to several in axillary and terminal glomerules. Calyx lobes triangular to lanceolate, 0.8-2 mm long, glabrous or minutely papillose. Corollas rotate, white, pink or lavender, very small and inconspicuous, to ca 0.7-1 mm long. Anthers 0.1-0.3 mm long, included. Style and stigma less than 0.5 mm long; stigmas included; styles thickened. Mature capsules 1.5-3 mm long, 1.5-2.5 mm wide, subglobose or slightly longer than wide, fully or 7/8 inferior, verrucose or papillose with minute rounded papillae, varying to glabrate. Seeds numerous per capsule, 0.1-0.3 mm long, trigonous. Flowering April to November. Chromosome number: 2n=36 (Lewis 1962).
Habitats: Wet or moist places, stream banks, lake shores, roadsides, ditches, fields, woods, savannahs, disturbed open places. Distribution: United States: Mainly in Atlantic and Gulf Coastal Plains and Mississippi Embayment in the southeastern states, also in other physiographic provinces, including Ozarks; southeastern Virginia, North Carolina (one county in southeast), South Car- olina (incl. Pickens Co.), southwestern (and one county in northern) Geor- gia, northern Florida, Alabama, Mississippi, central and western Tennessee, Louisiana, Arkansas, southeastern Missouri, eastern Oklahoma and eastern Texas.
Oldenlandia bosci does not seem to vary much in morphology over its range, in contrast to most other taxa in this and related genera.
4. Oldenlandia salzmanniu (DC.) Benth. & Hook. f. ex B.D. Jacks., Index Kew. 2:336. 1894; Benth. & Hook. f., Gen. Pl. 2:58. 1873. ined. (see Fosberg & Terrell 1985). Anotis salzmanni DC., Prodr. 4:433. 1830. TYPE: “Bahiam,” Salzmann s.n. (not seen). Hedyotis salzmannu (DC.) Steud., Nomencl. Bot., ed. 2. 1:726. 1840.
Hedyotis thestifolia A. St. Hil., Voy. Distr. Diam. 1:397. 1833. Olden- landia thesttfolia (A. St. Hil.) K. Schum. in Martius, Fl. Bras. 6, 6:270, pl. 127, f. 1. 1889.
Small creeping perennial herb. Stems slender, prostrate, rooting at nodes, glabrous. Leaves sessile or subsessile, broadly elliptic to ovate, 1.5-5.2 mm long, 0.7-2.5 mm wide, glabrous to sparsely hirsute. Stipules to ca 0.5 mm long, entire or with 1 to few marginal white hairs to 0.7 mm long. Flowers heterostylous, solitary, terminal and axillary on slender pedicels 3-12 mm long. Calyx lobes lanceolate, ovate, or oblong, 1-2.2 mm long, glabrate. Corollas subsalverform, pink, lavender, purple or white, 2.5-5 mm long, tube 1-2.2 mm
Terrell: Synopsis of Oldenlandia in United States 131
long, short and broad, densely hirsutulous or pubescent within: lobes ovate, equalling or longer than the tube. Anthers 0.5-0.8 mm long. Stigma branches (0.5-)1-1.5 mm long. Pin flowers with stigmas exserted 1-2 mm and anthers at distal end of tube; thrum flowers with anthers exserted on filaments | mm long and the stigmas included. Mature capsules ca 1.5 mm long and wide, subglobose, 7/8 to fully inferior, hirsute. Seeds ca 4-14 per capsule, 0.3-0.5 mm in diameter, trigonous. Flowering June to August, at least in western Florida and adjacent Alabama. Chromosome number: n=15, 2n=30 (Lewis 1966a).
Habitats and distribution: South America: Brazil, Argentina. Uruguay, Paraguay. United States: Locally established, apparently accidentally, near Pensacola, Escambia Co., western Florida and in adjacent Baldwin Co., Al- abama, in roadside ditch, by ponds and at edge of a marsh, where discovered by J.R. Burkhalter of Pensacola (Fosberg & Terrell 1985). The name, O.
salzmannii, is here accepted as correct pending study of the type specimens.
5. Oldenlandia callitrichoides Griseb., Mem. Amer. Acad. Arts n.s. 8:506. 1863. TYPE: Discussed in pending publication, as noted below. Hedyotis callitrichordes (Griseb.) W.H. Lewis, Rhodora 63:222. 1961.
Small creeping perennial herb. Stems soft, thin, prostrate, rooting at nodes, glabrous. Leaves with slender petioles 0.5-3 mm long; blades thin, suborbicular, broadly ovate or subrhombic, 0.5-4 mm long, 0.5-4.8 mm wide, sparsely covered above with scattered stiff white hairs to ca 0.5 mm long or infrequently glabrate. Stipules minute, with 0 to few whitish marginal hairs. Flowers apparently homostylous, solitary from axils on filiform pedicels to ca 2 cm long. Calyx lobes lanceolate or ovate, 0.3-1 mm long, cilated along the margins and sinuses. Corollas subsalverform, white or lobes purplish tipped, 1.5-3.5 mm long, tube slightly shorter than lobes. Anthers 0.2-0.3 mm long, exserted at mouth of tube on filaments to 0.3 mm long. Style and stigma 1.5-2 mm long, with the stigma exserted to or beyond anthers. Mature capsules 1- 2.7 mm long, 0.5-2 mm wide, narrowly turbinate or narrowly obconic, widest at apex, 9/10 to fully inferior, glabrous below ciliate calyx lobes, eventually splitting apart into 4 narrow segments. Seeds ca 20-35 per capsule, 0.3-0.5 mm long, ellipsoid, rhomboid or obtusely several angulate. Flowering September to January in Bahamas (Correll 1982). Chromosome number: 2n=22 (Terrell, et al. 1986).
Habitats and distribution: Moist or wet soil in grassy places, disturbed places, on rocks and cliffs, lawns, walls, curbs, stonework. West Indies, Panama, Yucatan. Adventive in Guyana; Sierre Leone in Africa; United States, Miami, Florida, at two locations: Vizcaya Art Museum on cement fountains and water channels; shaded, grassy place at Fairchild Tropical Garden; Hawaii: Oahu, Maui. As noted in Introduction, this species is being placed in a new genus
by Terrell and W.H. Lewis.
132 PHYTOLOGTIA volume 68(2):125-133 February 1990
ACKNOWLEDGMENTS
I thank the curators of the following herbaria for loans or for arranging access to collections during visits; ALU, AUA, DUKE, DUR, FLAS, FSU, GA, KY, LAF, LSU, MO, NCU, NO, NY, OKL, OKLA, SMU, TENN, TEX, UNA, US, USCH, USF, UWFP and VPI. Drs. W.H. Lewis and J.L. Reveal helpfully reviewed the manuscript. This paper is Scientific Article No. A-5054, Contribution No. 8111, of the Maryland Agricultural Experiment Station.
LITERATURE CITED
Bremekamp, C.E.B. 1952. The African species of Oldenlandia L. sensu Hi- ern. et K. Schumann, Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 48:1-297.
Correll, D.S. & H.B. Correll. 1982. Flora of the Bahama Archipelago. J. Cramer, Vaduz. 1692 pp.
Fosberg, F.R. 1943. The Polynesian species of Hedyotis (Rubiaceae). Bernice P. Bishop Mus. Bull. 174. 102 pp.
& E.E. Terrell. 1985. A recently established exotic in west Florida and Alabama (Hedyotis salzmannw or Oldenlandia salzmannit; Rubi- aceae). Castanea 50:49-51.
Halle, N. 1966. Rubiacées, pt. 1. III. Hedyotidées. pp. 75-124, in A. Aubreville, editor. Flore du Gabon. Mus. Hist. Nat., Paris.
Hepper, F.N. & R.W.J. Keay. 1963. Rubiaceae (part), vol. 2, Oldenlandia. pp. 210-212, in F.N. Hepper, editor. Flora of West Tropical Africa, 2nd ed. Crown Agents, London.
Hitchcock, A.S. & M.L. Green. 1929. Standard-species of Linnaean genera of Phanerogamae. Intl. Bot. Congress, Cambridge, 1930. Nomencl. Prop. Brit. Bot. 110-199, London.
Lewis, W.H. 1959. Chromosomes of east Texas Hedyotis (Rubiaceae). Southw. Naturalist 3:204-207.
. 1961. Merger of the North American Houstonia and Oldenlandia under Hedyotis. Rhodora 63:216-223.
Terrell: Synopsis of Oldenlandia in United States 133
Lewis, W.H. 1962. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. J. Bot. 49:855-865.
1964. Oldenlandia corymbosa (Rubiaceae). Grana Palynolog. 5:330-341.
1965. Cytopalynological studies of African Hedyotideae (Rubi- aceae). Ann. Missouri Bot. Gard. 52:182-211.
. 1966a. Chromosome numbers of Phanerogams, 1. Ann. Missouri
Bot. Gard. 53:100-103.
1966b. Chromosome numbers of Oldenlandia corymbosa (Rubi- aceae) from southeastern Asia. Ann. Missouri Bot. Gard. 53:257-264.
Standley, P. 1918. Rubiaceae. Oldenlandieae, in, North American Flora 32, part 1:17-39.
Terrell, E.E. 1975. Relationships of Hedyotts fruttcosa L. to Houstonia L. and Oldenlandia L. Phytologia 31:418-424.
_______, W.H. Lewis, H. Robinson & J.W. Nowicke. 1986. Phylogenetic im- plications of diverse seed types, chromosome numbers, and pollen mor- phology in Houstonza (Rubiaceae). Amer. J. Bot. 73:103-115.
Verdcourt, B. 1976. Rubiaceae (Part 1), Oldenlandia. pp. 268-315, in, R.M. Polhill, editor. Flora of Tropical East Africa. Crown Agents, London.
Willis, J.C. (rev. by H.K. Airy Shaw). 1973. A Dictionary of the Flowering Plants and Ferns, 8th ed. Cambridge Univ. Press, Cambridge, England. 1245 pp.
Phytologia (February 1990) 68(2):134-136.
NEW NAMES AND COMBINATIONS IN MEXICAN ALLOJSPERMUM (ASTERACEAE: HELIANTHEAE)
B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
Six new specific combinations and two new varietal combinations for the genus Alloispermum are proposed, as follows: A. gonzalezae (B. Turner) B. Turner, A. longiradiatum (Urbatsch & B. Turner) B. Turner, A. michoacanum (B.L. Robins.) B. Turner, A. michoa- canum var. liebmannii (Schultz-Bip. ez Klatt) B. Turner, A. palmeri (S. Wats. ex A. Gray) Fernandez & Urbatsch, A. palmeri var. lanci- folium (Urbatsch & B. Turner) Fernandez & Urbatsch and A. trida- coides (Urbatsch & B. Turner) Fernandez & Urbatsch. All of these are transfers from the genus Sabazia.
KEY WORDS: Alloispermum, Asteraceae, Heliantheae, systema- tics, México.
As treated by Fernandez & Urbatsch (in manuscript), Allozspermum is a genus of perhaps 14 species, mostly confined to montane habitats of Mexico, Central America and northern South America. It is very closely related to Sabazia and may ultimately encompass that taxon. The two are kept apart largely on features of habit and capitulescence, Allotspermum having rather simple stems that arise from short, stout, subterranean rhizomes and a ca- pitulescence that arises from primary peduncles that branch near the summit forming cymules of 3 to numerous heads. Species of Sabazia, as treated by the present author, have annual taproots or, when perennial, aerial stems arise from slender, wiry, rhizomes, the lower stems often decumbent and rooting at the nodes; in addition, the heads are usually solitary on elongate terminal or axillary peduncles.
Urbatsch & Turner (1975) placed several of the species included here in the genus Sabaziza, having removed these from Calea where they were anomalous. Robinson (1978) subsequently resurrected the older generic name, Allozsper- mum, to house these and we accept the reality that they are properly posi- tioned there. McVaugh (1984), in his usual conservative vein, retained most of the aforementioned anomalies in his heterogeneous Calea, but also maintained
134
Turner: New names and combinations in Mexican Allotspermum 135
Sabazia (by insertion of the latter within Calea, he creates a phyletically inde- fensible position). In any case, I intend to retain Calea, Sabazia and Allozsper- mum in my treatment of the tribe Heliantheae for México, believing these to be separate, closely related phyletic lines. Additional work will be needed to resolve the relationship of the latter two genera with the closely related genera Galinsoga and Schistocarpha, as noted by Fernandez & Urbatsch.
Alloispermum gonzalezae (B. Turner) B. Turner, comb. nov. BASIONYM: Sabazia gonzalezae B. Turner, Phytologia 63:307. 1987.
As noted in my original description, inclusion of this taxon in Sabazia was provisional. A review of the entire Allotspermum-Sabazia-Galinsoga complex now leads me to believe that it is properly positioned in Alloispermum, near A. tridacozdes.
Alloispermum longiradiatum (Urbatsch & B. Turner) B. Turner, comb. nov. BASIONYM: Sabazia longiradiata Urbatsch & B. Turner, Brittonia 27:353. 1975.
This species is superficially similar to A. palmer: but differs in its shorter stature, fewer leaves along the stems and much longer rays. On habital features and its branched capitulescence it belongs to Allowspermum.
Alloispermum michoacanum (B.L. Robins.) B. Turner, comb. nov. BA- SIONYM: Sabazia michoacana B.L. Robins., Proc. Amer. Acad. Arts 27:173. 1892.
Two varieties are recognized under this species, as follows:
1. Peduncular hairs with purple crosswalls; leaves mostly widest at or near RUMEN ool op Nahmias) slanted miata ailaly aia ae eee ae var. liebmannit
1’ Peduncular hairs without purple crosswalls; leaves widest at or near the mites: Nichowedta io /iicbie los 2 ety ohne ves sae ees var. michoacanum
Alloispermum michoacanum var. michoacanum. Sabazia michoacana B.L. Robins., Proc. Amer. Acad. Arts 27:173. 1892. Sabazia liebmannii var. michoacana (B.L. Robins.) Longpre.
Known only from Michoacan in pine-oak forests, 2000-3100 m.
Alloispermum michoacanum var. liebmannii (Schultz-Bip. ez Klatt) B. Turner, comb. nov. BASIONYM: Sabazia hebmannw Schultz-Bip. ez Klatt, Leopoldina 23:90. 1887. Not Alloispermum hebmanniz (Schultz- Bip. ez Klatt) H. Robins., Phytologia 38:412. 1978, which is based upon Calea liebmannii Schultz-Bip. er Klatt, Leopoldina 23:145. 1887. Sabazia liebmannit Schultz-Bip. ez Klatt var. lebmanniz.
136 PHYTOLOGIA volume 68(2):134-136 February 1990
Sabazia liebmannit var. hintoni Longpre.
Sabazia hebmanni var. ovatifolia Longpre.
Mexico State, Guerrero and Oaxaca in pine-oak woodlands, 2000-2900 m. A variable taxon but readily distinguished from var. michoacanum by the characters given in the above couplet.
Alloispermum palmeri (S. Wats. er A. Gray) Fernandez & Urbatsch, comb. nov. BASIONYM: Calea palmerz S. Wats. ex A. Gray, Proc. Amer. Acad. Arts 22:430. 1887.
Alloispermum palmeri var. lancifolium (Urbatsch & B. Turner) Fernan- dez & Urbatsch, comb. nov. BASIONYM: Sabazia palmert var. lancifolia Urbatsch & B. Turner, Brittonia 27:353. 1975.
Alloispermum tridacoides (Urbatsch & B. Turner) Fernandez & Urbatsch, comb. nov. BASIONYM: Sabazia tridacoides Urbatsch & B. Turner, Brittonia 74:351. 1975.
ACKNOWLEDGMENTS
I am grateful to Dr. Guy Nesom and Dr. Andrew McDonald for reviewing the present paper and to Dr. Lowell Urbatsch for sending me an advance copy of his paper with Fernandez and publishing in advance their several new combinations which they will treat in more detail in their forthcoming systematic study of the genus.
LITERATURE CITED
Fernandez, C. & L. Urbatsch. (in manuscript). Systematics of the genus Alloispermum (Asteraceae-[leliantheae).
McVaugh, R. 1984. Caleain Flora Novo-Galiciana 12:187-200.
Robinson, H. 1978. Restoration of the genus Alloispermum. Phytologia 38:411-412.
Urbatsch, L. & B.L. Turner. 1975. New species and combinations in Sabazia AHeliantheae, Galinsoginae). Brittonia 27:348-354.
Phytologia (February 1990) 68(2):137.
CORRECTIONS IN THE TYPE LOCALITY CITATION OF VERBESINA HOWARDIANA
John Olsen Department of Biology, Rhodes College, Memphis, Tennessee 38112 U.S.A.
ABSTRACT
The type locality of Verbesina howardiana was incorrectly cited as the Dominican Republic. The specimen cited was actually collected on the island of Dominica.
KEY WORDS: Verbesina, Asteraceae, Lesser Antilles, systematics.
In the paper containing the original description of Verbesina howardiana (Olsen 1989), the species was cited to be found in the Dominican Republic. In fact, the plant has been found only on the island of Dominica. The error resulted from a mistake in my concept of Caribbean geography, as an incorrect connection was made between Dominica (where the specimens originated) and “the Dominican” (a common reference to the Dominican Republic).
Corrections are required in three places in the original publication, all on page 107. The first is in the title, where DOMINICA should replace THE DOMINICAN REPUBLIC. Second, in the abstract, Dominica should replace Hispanola. The third error is in the type citation where DOMINICA should replace DOMINICAN REPUBLIC.
ACKNOWLEDGMENTS
I wish to thank Dr. Dan Nicolson for bringing the error to my attention.
LITERATURE CITED
Olsen, J. 1989. A new species of Verbesina section Verbesinaria from the
Dominican Republic. Phytologia 67:107-108.
137
Phytologia (February 1990) 68(2):138-140.
A NEW SPECIES OF APHANACTIS (ASTERACEAE: HELIANTHEAE; GALINSOGINAE), FROM OAXACA, MEXICO
B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.
ABSTRACT
A new species, Aphanactis macdonaldii B. Turner, from Cerro Quiexobra, 3650-3800 m, Oaxaca, México is described and illustrated. The genus was previously unknown to México. The species is notable for its large heads with bright yellow rays, unlike any previously described
species.
KEY WORDS: Aphanactis, Asteraceae, Heliantheae, systematics, México, Oaxaca.
Routine identification of a remarkable assemblage of species belonging to the Asteraceae collected by Dr. Andrew McDonald from the subalpine region of Cerro Quiexobra (3650-3800 m) resulted in the description of seven new species from that peak alone (Nesom, in prep.; Turner, in prep.). The most remarkable discovery has been the present species, which is placed in Aphanactis, the first report for that genus from México, although two species are known from Central America (Turner 1980).
Aphanactis macdonaldii B. Turner, sp. nov., Figure 1. TYPE: MEXICO. Oaxaca: 35 km ESE of Miahuatlan, 5 km NE of Santo Domingo Ozolote- pec, Cerro Quiexobra and vicinity, 3650-3800 m (16° 10’ N, 96° 15’ W), timberline vegetation in open glades along ridges and in mountain sad- dles, 10 Dec 1989, Andrew McDonald 2904 (HOLOTYPE: TEX; Isotype: MEXU).
Aphanactis obtusatae (S.F. Blake) B. Turner similis sed foliis
majoribus plerumque basalibus et capitulis majoribus ligulis luteis profunde 3-lobatis differt.
138
139
, México
New Aphanactis from Oaxaca
A om
ctis macdonaldii, from holotype.
Aphana
ria, 7.
140 PHYTOLOGIA volume 68(2):138-140 February 1990
Perennial, nearly scapose, herbs 15-25 cm high, the stems erect and aris- ing from relatively thick lateral rhizomes. Lowermost leaves mostly opposite, sessile, linear-oblanceolate, much reduced and remote along the upper portion of the stem, the blades with 3-7 nearly parallel nerves, moderately hirsutu- lous throughout, the apices 3 lobed or not. Heads hemispheric, ca 3 cm wide across the extended rays, borne singly on nearly leafless scapes 5-20 cm long. Involucres 9-11 mm high, the bracts ovate-elliptical, mostly subequal in 2-3 series, pilose like the peduncles. Receptacle broadly conical, the chaff filiform. Ray florets 11-13, pistillate, fertile, the ligules bright yellow, deeply 3 lobed, ca 15 mm long, 7-9 mm wide, the tube ca 2.5 mm long, pubescent. Disk florets numerous, the corollas yellow, the tube pubescent, ca 1 mm long, the limb glabrous, ca 2 mm long with 5 lobes ca 0.5 mm long. Achenes ca 3 mm long, obovate, 5 ribbed, black, glabrous, epappose.
The species is remarkable for its large heads with bright yellow rays. In- deed, it is so distinct from the other species in habit that I originally thought it might belong to Selloa (which it superficially resembles) or else might rep- resent an undescribed genus. Detailed comparison of the heads, florets and achenes with those of Aphanactis, however, strongly suggests that it goes with or near that genus. So positioned, the characters of the genus are expanded to include large, yellow, 3 lobed ligules (vs single and white or pinkish) and disk florets with rather tubular throats.
It is a pleasure to name this remarkable species for its remarkable collector, Dr. Andrew McDonald, avid student of the alpine vegetation of Mexico and expert on the family Convolvulaceae.
ACKNOWLEDGMENTS
I am grateful to Dr. Guy Nesom for the Latin diagnosis and to him and Dr. McDonald for a review of the manuscript itself. Ms. Nancy Webber provided the illustration.
LITERATURE CITED
Turner, B.L. 1980. La taxonomia del genero Aphanactis (Asteraceae - He- liantheae). Bol. Soc. Argentina Bot. 19:33-44.
Phytologia (February 1990) 68(2):141-143.
PANICUM RIGIDULUM VAR. COMBSII (POACEAE) IN LOUISIANA
Paul M. McKenzie’, Lowell E. Urbatsch? & Michel G. Lelong’ 'School of Forestry, Wildlife and Fisheries, Louisiana Agricultural Experiment Station, Louisiana State Agricultural Center, “Department of Botany, Louisiana State University, '“Baton Rouge, Louisiana 70803 & ‘Department of Biology, University of South Alabama, Mobile, Alabama 36688 U.S.A.
ABSTRACT
The occurrence of Panicum rigidulum var. combsi is documented in Louisiana from Natchitoches Parish.
KEY WORDS: Floristics, Panicum, Poaceae, Louisiana.
Hitchcock & Chase (1910) listed seven species for the Agrostoidea group of Panicum: P. agrostoides Spreng., P. anceps Michx., P. combsz Scribn. & Ball, P. condensum Nash., P. longifolium Torr., P. rhizomatum Hitchce. and P. stipitatum Nash. Fernald (1934; 1950) listed P. combsz as a variety of P. longifolium: P. longifolium Torrey var. combsit (Scribn. & Ball) Fern. Within the group Agrostoidea, Hitchcock (1951) included the same seven species as in Hitchcock’s & Chase’s (1910) treatment, but also included P. abscissum Swallen, and retained P. combsii at the species level. Gould (1975) considered P. agrostoides, P. condensum, P. longifolium and P. stipitatum to be synony- mous with P. rigzdulum Bosc ez Nees. Lelong (1984) retained varietal status for P. combsw within P. rigidulum.
Panicum rigidulum var. combsii (Scribn. & Ball) Lelong is not listed for Louisiana by Allen (1980) and MacRoberts (1988) found no reports of it for Louisiana. Based on a specimen taken at Lake Charles by Chase (Chase 4434, US), Hitchcock & Chase (1910) and Hitchcock (1951) included Louisiana within the range of this taxon treated as P. combsi. Additionally, Fernald (1934) listed a specimen of the taxon from New Orleans, but no information was provided on the collector, collection number, or the herbarium where the specimen was located.
A recent specimen of P. rigidulum var. combsti for Louisiana was collected in Natchitoches Parish, in dry, sandy soil of a longleaf pine (P. palustris Mill.) hillside, above a hillside seep, northwest of the intersection of Middle Branch Rd. and U.S. Forest Service Rd. 321, 2 Aug 1985 (McKenzte 171 with L.E. Urbatsch, Annette Parker and Karla Wilzer, LSU; annotated by M. Lelong,
141
142 PUY Tay Ore fA volume 68(2):141-143 February 1990
March 1989). This site is more similar to the “wet pine-barrens” designated for the taxon by Fernald (1950) than to the “marshes, shores of lakes and ponds” described as the habitat in Mississippi (Lelong 1986). The paucity of collections from Louisiana suggests that P. rigzdulum var. combsizis rare in the state. It is likewise rare in Mississippi, where it is known only from Harrison County (Lelong 1986).
Within the typical subgenus of Panicum, P. rigidulum is recognizable by its short knotty bases or caudexes, often densely tufted culms, and strongly com- pressed sheaths and culms. Panicum rigidulum var. combs differs from the more widespread and common typical variety by its narrower more pubescent leaves and larger spikelets. The former variety possesses adaxially pilose leaf blades usually 2-7 mm wide with fimbriate-ciliate membranous ligules, spikelets 2.6-3.7 mm long and usually purple, slender, erect pedicels.
ACKNOWLEDGMENTS
We thank Mary Sangrey, U.S. National Herbarium, Washington, D.C. and the Department of Botany, Louisiana State University, Baton Rouge for their assistance with this report.
LITERATURE CITED
Allen, C 1980. The Grasses of Louisiana. Southwestern Louisiana Press, Lafayette, 358 pp.
Fernald, M.L. 1934. Realignments in the genus Panicum. Rhodora 36:61-87.
______. 1950. Gray’s Manual of Botany, 8th ed. American Book Co., New York, 1632 pp.
Gould, F.W. 1975. The Grasses of Teras. Texas A&M Univ. Press, College Station, 653 pp.
Hitchcock, A.S. 1915. Afanual of the ae of the United States, 2nd ed. revised by A. Chase. U.S.D.A. Misc. Publ. no. 200, 1051 pp.
& A. Chase. 1910. The North American species of Panicum. Contr. U.S. Natl. Herb. 15:1-396.
Lelong, M.G. 1984. New combinations for Panicum subgenus Panicum and subgenus Dicanthelium (Poaceae) of the southeastern United States.
Brittonia 36(3):262-273.
McKenzie, et al.: Panicum rigidulum var. combsu in Louisiana 143
Lelong, M.G. 1986. A taxonomic treatment of the genus Panicum (Poaceae)
in Mississippi. Phytologia 61(4):251-269.
MacRoberts, D.T. 1988. A Documented Checklist and Atlas of the Vascular Flora of Louisiana. Part 1. Pteridophyta, Gymnospermae, and Mono- cotyledoneae. Louisiana State University in Shreveport, 256 pp.
Phytologia (February 1990) 68(2):144-155.
TAXONOMIC SUMMARY OF ERICAMERIA (ASTERACEAE: ASTEREAE), WITH THE INCLUSION OF HAPLOPAPPUS SECTS. MACRONEMA AND ASIRIS
Guy L. Nesom Department of Botany, University of Texas, Austin Texas 78713 U.S.A.
ABSTRACT
Ericameria sensu stricto (12 species) is broadened to include the species of sect. Stenotopsis (1 species), sect. Macronema (Nutt.) Ne- som (9 species) and sect. Asiris (H.M. Hall) Nesom (5 species). The nomenclature for the 27 species of Ericameria as so defined is sum- marized and criteria for the distinction of the sections are presented in a key. New specific combinations are proposed for E. compacta, E. crispa, E. discoidea, E. gilmanii, E. greenei, E. obovata, E. ophitidis, E. suffruticosa, E. watsonii and E. zionis. One new
varietal combination is proposed, Ericameria discoidea var. linearis. KEY WORDS: Ericameria, Haplopappus, Asteraceae, Astereae.
Ericameria has been understood to include a group of subshrubby species with narrow, entire, punctate-resinous leaves, small heads commonly in corym- boid capitulescences and a base chromosome number of z=9. The genus has not been generally accepted by floristicians since Hall’s treatment of it as Haplopappus sect. Ericameria (Hall 1928), although recent studies (primarily Johnston 1970 and Urbatsch, 1975; 1976; 1978; 1979) have recognized it as distinct.
Nesom, et al. (submitted) have sharpened the definition of Ericameria by removing from it as a separate genus, seven species (see Excluded Species, be- low) closely related to Euthamza, but distantly related to species traditionally recognized as Haplopappus. Plants of the new genus can be distinguished mor- phologically by characteristics of their involucral bracts, which have a white indurated, enervate basal portion and a glandular herbaceous patch on the up- per portion, and their disc corollas, which are zygomorphic, the lobes strongly uneven in length. In contrast, plants of Ericameria have involucral bracts without an apical glandular patch, but with a clear midvein from base to tip and their disc corollas are regular with lobes of even length.
144
Nesom: Taxonomic summary of Ericameria 145
Although Hall (1928) segregated the species of Haplopappus sect. Asiris within his broad concept of Haplopappus, some were originally included in Eri- camerza by Nuttall, and some were again included in that genus by Urbatsch in his contribution to a checklist of North American plants (Kartesz & Kartesz 1980). One of the six species originally recognized by Hall in sect. Aszris, H. purpusii, is included in the new genus being described by Nesom, et al. (sub- mitted). A connection between Macronema (Haplopappus sect. Macronema) and Ericameria has not been generally recognized, although Macbride (1918, see comments below) transferred Haplopappus bloomeri to Ericameria. A close similarity between sect. Macronema and sect. Asitris, however, has been ac- knowledged as they are treated together in keys (e.g., Ferris 1960; Cronquist 1973). Ericameria linearifolia, one of the two species of Stenotopsis ( Hap- lopappus sect. Stenotopsis), has been transferred to Ericameria by Urbatsch & Wussow (1979). The other species, E. parrasana, is part of the new genus.
An overview of the taxonomy of Ericameria has not been published since Hall’s treatment of Haplopappus. In the course of studying the generic bound- aries of Ericamerza for a floristic treatment of the Mexican species as well as for the separation of a new genus, I have arrived at a broadened view of the former, which is presented below.
There are two primary areas of difficulty in formulating a clear definition of Ericamerza, the first involving sect. Stenotopsis, the second involving sects. Macronema and Asirts.
Section Stenotopsis
The first problem involves the relationship of typical Ericameria with E. lineartfolia, which was segregated as the genus Stenotopsis Rydb. and included as one of the two species of Haplopappus sect. Stenotopsis (Rydb.) H.M. Hall. Extensive and well documented natural hybridization (Urbatsch & Wussow 1979; Cody & Thompson 1986) exists between £. linearifolia, which has long, merely bracteate peduncles with large, solitary heads with long, prominent ray flowers and 3 veined, stipitate glandular phyllaries, and EF. cooper, which is morphologically more typical of Ericameria. These two species have similarly colored pappus bristles and similarly shaped style appendages, and because of this, they are considered by Urbatsch & Wussow to be closely related and both placed in Ericameria sect. Stenotopsis. Considering the large differences be- tween these two species, however, the small morphological similarities used by these workers to unite them are likely to be fortuitous. If the main criterion for associating the two species is ease of hybridization, attempts of artificial crosses between E. linearifolia and other species of Ericameria should be considered in the formulation of more meaningful hypotheses of close relationship. This is particularly true in view of the natural hybrids known between Haplopappus macronema and Chrysothamnus nauseosus (Anderson & Reveal 1966), which also are extremely divergent in morphology.
146 PHY TOLOGTIA volume 68(2):144-155 February 1990
Although Ericamerza linearifolza falls outside the boundaries of typical Ert- cameria in some features, it produces somewhat flattened, 6-8 nerved achenes and punctate leaves, which are characteristic of the genus. Its large, soli- tary heads and long ray flowers are more similar to those of species of sect. Macronema. Ericameria coopert is much more similar to typical Ericamerza in its small, discoid, apically clustered heads, but its turbinate-subcylindric achenes with 10-12 thin nerves are atypical.
Sections Afacronema and Asiris
The second problem in defining Ericameria involves its distinction from Haplopappus sects. Macronema and Asiris. The following key provides con- trasts that, with the caveats discussed below, separate these groups from Erv- camerta and Stenotopsis.
1. Leaves flat and obovate to terete and linear, sometimes in axillary fas- cicles, usually resinous from punctate glands; involucral bracts with a thick to thin, prominent, orange resinous midvein often expanded at the very apex, the bract apices rounded to acute but apiculate or ap- pendaged in 2 species; collecting appendages of the disc style branches most narrowly triangular and equal or shorter in length than the stig- matic portions, rarely linear and longer; achenes narrowly oblong, most commonly compressed or flattened, with (4-)6-8(-12) nerves, sometimes Oe RS. Uae ee ee ee ee RAEI 2
2. Heads mostly in panicles or corymboid capitulescences, solitary in one species; phyllaries 1 nerved, papillate glandular in one species but not stipitate glandular; ray flowers absent or with short, incon- a Ee Re ney ae a a ee sect. Ericameria
2’ Heads solitary; phyllaries 3 nerved, stipitate glandular; ray flowers with long prominent lipules 00's). 68. 08. OS sect. Stenotopsis
1’ Leaves narrow and mostly flat, not in axillary fascicles, resinous but ap- parently eglandular or with stipitate glands; midvein of the involucral bracts orange resinous to greenish yellow, the bract apices rounded to acute, with a pronounced, herbaceous apiculum or appendage; collecting appendages of the disc style branches linear to linear lanceolate, longer than the stigmatic portions; achenes narrowly cylindric to flattened, 3-5 wien bi wed! ecru saythe ta. ahaa sai 4 nid: bck am cil ‘den iy-nss inet anal 3
3. Heads relatively large, solitary to clustered, immediately subtended by leaf like bracts; involucral bracts apically apiculate to appendaged, with a definite, orange resinous midvein, not keeled; achenes cylin- dric to slightly compressed, 5 nerved ............ sect. Macronema
Nesom: Taxonomic summary of Ericameria 147
3’ Heads relatively small, clustered, without definite leaf like bracts; involucral bracts apically apiculate, with a thin, greenish yellow midvein, often slightly keeled; achenes distinctly flattened, 3-4 OPE Ps acta ce cal et os care cain ane tee ee sect. Asiris
Ericameria, Macronema and Asiris each comprise species with variably shaped leaves and solitary to clustered heads variable in size. The heads in both may be eradiate or radiate with ray corollas variable in size. The distinction between them appears to lie in the nature of the leaf glandularity, the shape and nervation of the achenes and the shape of the style branch collecting appendages and their length relative to the stigmatic portion. The species of sect. Asir1s and those of sect. Afacronema are more closely similar between themselves, as evidenced by their apiculate involucral bracts, long linear style appendages and few nerved achenes. The species of sect. Aszris are divergent from the species of Macronema in their narrower heads with more thinly herbaceous involucral bracts and their peduncles that are not so strongly leafy.
The definition, however, between Ericamerza and Macronema loses signif- icant clarity because of overlapping variation in both groups. This is particu- larly true in Ericamerza, where E. pinifolia is strongly similar to Macronemain its apiculate to appendaged involucral bracts, linear style branches longer than the stigmatic portions and narrowly cylindrical achenes. Further, the leaves of some plants of this species are not at all punctate. The generic affinity of EF. pintfolia has never been questioned and it belongs firmly in Ericameria, where it is closely related to the type species of the genus, E. ertcotdes, which also shows some of the same Macronema like features. The differences between the groups in achene shape and nervation are not constant, because FE. palmerz has achenes typical of Macronema, terete with (4-)5(-7) nerves. The achenes of E. cooper: are terete to slightly compressed, and along with those of several other species, FE. partshit and E. pinifolia, may produce up to 12 nerves.
Finally, some plants of Haplopappus (sect. Macronema) bloomerz produce leaves that clearly are punctate resinous. Macbride’s (1918) transferral of this species to Ericameria was made without specific comment, but perhaps re- flected his reliance upon this criterion to distinguish Ericameria. Haplopappus bloomerz is highly variable in a number of other characteristics, as evidenced by the number of infraspecific taxa that have been named within it (see Hall 1928). Ericameria pintfolia is equally as variable, and because the variabil- ity in each species includes forms that are morphologically “shifted” toward the other, an investigation of these species for the possibility of hybridization should be interesting.
Parallel variation in Chrysothamnus
Chrysothamnus appears to be very closely related to Ericameriaand Macro- nema, particularly the latter (Anderson 1970), but it is generally accepted as a
148 PABYY) TOLL OoG PA volume 68(2):144-155 February 1990
distinct genus (Hall & Clements 1923; Blake 1926; Anderson 1984). It is more homogenous than either Ericameria or Macronema in its densely arranged, narrow, strictly eradiate heads and its involucral bracts in vertical files. Sub- stantial variation occurs within Chrysothamnus, however, in the same charac- ters that separate Ericameria from ‘\facronema (Anderson 1970). The leaves of most species are resinous but non punctate, yet they are punctate in oth- ers. The achenes are variable in shape (terete to flattened) and in number of nerves, and the style branch collecting appendages vary from shorter to longer than the stigmatic portions. The species have been arranged into sections by Anderson (1984) to account for aspects of this variability.
Other z=9 groups of Haplopappus: Hesperodoria, Petradoria, Stenotus, Tonestus and Oreochrysum
Hesperodoria E. Greene ( Haplopappus sect. Hesperodoria |E. Greene] H.M. Hall), with slightly resinous punctate leaves, may be related to the group of genera around Ericameria, but its scabrous margined leaves and strongly turbinate heads are unlike any species there. In its general habit, it is more like Petradoria, whose composition and systematic position has been some- what ambiguous, although it appears to be closely related to Chrysothamnus (Anderson 1963; 1983; 1984).
Stenotus Nutt. (Haplopappus sect. Stenotus [Nutt.] A. Gray) appears to be situated outside of the closely related elements of the Ericameria group, contrary to an earlier hypothesis (Nesom 1989). In contrast to Ericameria and Macronema, as well as Chrysothamnus (excluding Petradoria), plants of Stenotus are uniformly low, caespitose and monocephalous herbs. They are perennials, but on the basis of morphology, Stenotus clearly does not belong in the Ertcamerza-Asiris-Macronema lineage as a “woody shrub” as indicated by Clark, et al. (1980), although it is similar in flavonoids to those taxa.
Plants of Tonestus A. Nels. (Haplopappus sect. Tonestus {A. Nels.] H.M. Hall) are also herbaceous and they are further characterized by thick caudex branches or rhizomes, plants mostly single stemmed from the base, leaves with a strong tendency to produce spinulose toothed margins and thin herbaceous bracts that nearly enclose the heads. Some of the species of Stenotus have been confused with Tonestus but the latter is clearly not a member of the Ericameria-Macronema alliance (Nesom & Morgan, submitted). Plants of the monotypic Oreochrysum Rydb. (Haplopappus sect. Oreochrysum [Rydb.| H.M. Hall) are rhizomatous, non resinous herbs with broad, relatively thin, clasping leaves and herbaceous, reflexing involucral bracts and could only be distantly related to Ericameria. Anderson & Creech (1975) included it within Sohdago. Apart from Ericameria and its close relatives as recognized in the present paper, and from Hesperodoria and Petradoria, the species of Stenotus, Tonestus and Oreochrysum are the others of Haplopappus (sensu Hall 1928) with a base chromosome number of z=9.
Nesom: Taxonomic summary of Ericameria 149
In summary, the species of Ericameria (12), Stenotopsis (1), Mfacronema (9) and sect. Aszris (5) constitute four apparently closely related lineages that are overlapping in morphology. Natural hybridization occurs between Ericameria and Stenotopsis. On morphological grounds, Chrysothamnus is also closely related to these groups and natural hybrids are known between Macronema and Chrysothamnus, but Chrysothamnus is generally accepted as a distinct genus. To provide a taxonomic framework for these four sections of Haplopappus sensu Hall that are closely related to Chrysothamnus, there are several options. First, Ericameria, Stenotopsis, Macronema and Asiris might each be recognized as a separate genus, or Ericameria (with Stenotopsis) and Macronema (with Asiris) could be recognized, but in either case, there would be no morphological features to consistently separate the generic units. Alter- natively, Ericameria could be expanded to bring the species of all four sections into a single taxon of coordinate rank with Chrysothamnus, resulting in the recognition of two closely related genera with similar patterns of variation among their respective species.
If, as hypothesized by Clark, et al. (1980) on the basis of flavonoid pro- files, Ericameria proves to be closest to the ancestral form in this group, with Macronema and perhaps Aszris as derivatives, and if the closest relative of Macronema proves to be Chrysothamnus, strict adherence to principles of cladistic classification would necessitate the merger of Chrysothamnus with all the rest. This would be extremely difficult to justify on a pragmatic basis, however, in view of the careful and detailed morphological and anatomical in- vestigations of Chrysothamnus by Loran Anderson, which have not suggested that it is congeneric with Macronema.
In order to clarify the boundaries of Ericameria, seven species have been removed as a separate, distantly related genus (Nesom, et al. submitted). In a correlated step, I propose to enlarge Ericameria, recognizing it as closely related to Chrysothamnus, and leaving as Haplopappus and its close relatives a group of species of South America (and North America if Hazardia is included) with the base chromosome number of z=5 (Brown & Clark 1982).
Taxonomic Summary of Ericameria
Ericameria Nutt., Trans. Amer. Philos. Soc., ser. 2 7:318. 1841. TYPE SPECIES: Ertcamerta microphylla Nutt., nom. nov. illeg. (= E. eri- cotdes).
As pointed out by Hall (1928), Nuttall arbitrarily adopted a new epithet (“microphylla”) when he transferred the type species to the new genus Eri- camerza. He cited “Haplopappus ericoides (Less.) DC.” as the name his new one would replace, but that combination was first made by Hooker & Arnott.
The following species are included, with partial synonymy.
150
PHY TOLOGIA volume 68(2):144-155 February 1990
A. Ericameria sect. Ericameria
ky
Ericameria arborescens (A. Gray) E. Greene, Alan. Bot. S.F. Bay Reg. 175. 1894. Bigelovia arborescens A. Gray, Proc. Amer. Acad. Arts 8:640. 1873. Haplopappus arborescens (A. Gray) H.M. Hall, Univ. California Publ. Bot. 7:273. 1919.
2. Ericameria brachylepis (A. Gray) H.M. Hall, Univ. California Publ. Bot.
si.
3b.
4a.
4b.
3:56. 1907. Bigelovia brachylepis A. Gray, Bot. California 1:614. 1876. Haplopappus brachylepis (A. Gray) H.M. Hall, Univ. California Publ. Bot. 7:273. 1919; non Phil. Haplopappus propinquus S.F. Blake, nom. nov., Contr. U.S. Natl. Herb. 23:1490. 1926.
Ericamerta cooperi(A. Gray) H.M. Hall, Univ. California Publ. Bot. 3:56.
1907. Bigelovia coopert A. Gray, Proc. Amer. Acad. Arts 8:640. 1873. Haplopappus cooperi (A. Gray) H.M. Hall, Carnegie Inst. Washington, Publ. 389:275. 1928.
Ericameria monactis (A. Gray) McClatchie, Erythea 2:124. 1894. Hap- lopappus monactis A. Gray, Proc. Amer. Acad. Arts 19:1. 1883.
Ericameria coopert var. bajacalifornica (Urbatsch & Wussow) Urbatsch, Phytologia 67:109. 1989. Ericameria cooperz subsp. bajacalifornica Ur- batsch & Wussow, Brittonia 31:274. 1979.
Ericameria cuneata (A. Gray) McClatchie, Erythea 2:124. 1894. Hap- lopappus cuneatus A. Gray, Proc. Amer. Acad. Arts 8:635. 1873.
Ericameria cuneata var. macrocephala Urbatsch, Madrono 23:344. 1976.
. Ericameria cuneata car. spathulata (A. Gray) H.M. Hall, Univ. California
Publ. Bot. 3:52. 1907. Bigelovia spathulata A. Gray, Proc. Amer. Acad. Arts 11:74. 1876. Haplopappus cuneatus var. spathulatus (A. Gray) S.F. Blake, Contr. U.S. Natl. Herb. 23:1849. 1926.
Ericameria ericoides (Less.) Jepson, Fl. W. Mid. Calif. 559. 1901. Diplopappus ericoides Less., Linnaea 6:117. 1831. Haplopappus ericoides (Less.) Hook. & Arn., Bot. Beechey Voy. 146. 1833; non DC., Prodr. 5:346. 1836. Ericameria microphylla Nutt., nom. illeg., Trans. Amer. Philos. Soc., ser. 2 7:319. i841.
6. Ericameria fasciculata (Eastw.) Macbr., Contr. Gray Herb. 56:36. 1918.
Chrysoma fasciculata Eastw., Bull. Torrey Bot. Club 32:215. 1905. Hap- lopappus eastwoodae H.M. Hall, nom. nov., Carnegie Inst. Washington,
Publ. 389:258. 1928.
Nesom: Taxonomic summary of Ericameria 151
7. Ericameria juarezensis (R. Moran) Urbatsch, Phytologia 67:109. 1989. Haplopappus juarezensis R. Moran, Trans. San Diego Soc. Nat. Hist. 15:154-155. 1969.
8. Ericameria larictfolia (A. Gray) Shinners, Field & Lab. 18:27. 1950. Haplopappus laricifolius A. Gray, Pl. Wright. 2:80. 1853.
Ericameria nelsoni (Fernald) S.F. Blake, Contr. Gray Herb. 52:26. 1917. Bigelovia nelsoni Fernald, Proc. Amer. Acad. Arts 36:505. 1901.
9. Ericameria martirensts Wiggins, Contr. Dudley Herb. 1:177. 1933. Aplopappus martirensis (Wiggins) S.F. Blake, Proc. Biol. Soc. Wash- ington 48:173. 1935.
10a. Ericameria palmert (A. Gray) H.M. Hall, Univ. California Publ. Bot. 3:53. 1907. Haplopappus palmerz A. Gray, Proc. Amer. Acad. Arts 11:74. 1876.
10b. Ericameria palmeri var. pachylepis (H.M. Hall) Nesom, Phytologia 67:104. 1989. Haplopappus palmerisubsp. pachylepis H.M. Hall, Carnegie Inst. Washington, Publ. 389:267. 1928.
lla. Ericameria parishu (E. Greene) H.M. Hall, Univ. California Publ. Bot. 3:55. 1907. Bzgelovia parish E. Greene, Bull. Torrey Bot. Club 9:62. 1882. Haplopappus parishit (E. Greene) S.F. Blake, Contr. U.S. Natl. Herb. 23:1491. 1926.
llb. Ericameria parishw var. peninsularts (R. Moran) Nesom, Phytologia 67:104. 1989. Haplopappus arborescens subsp. peninsularis R. Moran, Trans. San Diego Soc. Nat. Hist. 15:152. 1969.
12. Ericameria pinifolia (A. Gray) H.M. Hall, Univ. California Publ. Bot. 3:54. 1907. Haplopappus pinifoltus A. Gray, Proc. Amer. Acad. Arts 8:636. 1873.
B. Ericameria sect. Stenotops1s (Rydb.) Urbatsch & Wussow, Brittonia 31:273. 1979. Stenotopsis Rydb., Bull. Torrey Bot. Club 23:617. 1900. TYPE SPECIES: Haplopappus lineartfolius DC. (= Ericameria linearn- folia [DC.] Urbatsch & Wussow). Haplopappus sect. Stenotopsis (Rydb.) H.M. Hall, Carnegie Inst. Washington, Publ. 389:156. 1928, in part.
1. Ericamerza linearifolia (DC.) Urbatsch & Wussow, Brittonia 31:273. 1979. Haplopappus linearifolius DC., Prodr. 5:347. 1836. Stenotus linearzfolius (DC.) Torrey & A. Gray, Fl. N. Amer. 2:238. 1842. Stenotopsis linear- tfoltus (DC.) Rydb., Bull. Torrey Bot. Club 27:617. 1900.
152 PHYTOLOGIA volume 68(2):144-155 February 1990
Haplopappus interior Coville, Proc. Biol. Soc. Washington 7:65. 1892. Haplopappus linearifolius var. interior (Coville) Jones, Proc. Cali-
fornia Acad., ser. 2 5:697. 1895.
C. Ericameria sect. Asiris (H.M. Hall) Nesom, comb. nov. BASIONYM: Haplopappus sect. Asiris H.M. Hall, Carnegie Inst. Washington, Yearb. 25:342. 1926. TYPE SPECIES: Ericamerza nana Nutt.
1. Ericameria cervina (S. Wats.) Rydb., Fl. Rocky Mts. 853. 1917. Hap- lopappus cervinus S. Wats., Amer. Naturalist 7:301. 1873.
2. Ericameria nana Nutt., Trans. Amer. Philos. Soc., ser. 2 7:319. 1841. Haplopappus nanus (Nutt.) D.C. Eaton, Bot. King’s Ezpl. 159. 1871. Chrysothamnus nanus (Nutt.) J.T. Howell, Fl. N.W. Amer. 302. 1900.
3. Ericameria obovata (Rydb.) Nesom, comb. nov. BASIONYM: Macrone- ma obovatum Rydb., Bull. Torrey Bot. Club 27:618. 1900. Haplopappus rydbergit S.F. Blake, nom. nov., Contr. U.S. Natl. Herb. 25:545. 1925; not Haplopappus obovatus Phil. Haplopappus watsoni var. rydbergi(S.F. Blake) S.L. Welsh, Great Basin Nat. 43:295. 1983. I have not been able
to evaluate Welsh’s taxonomic judgment.
4. Ericameria resinosa Nutt., Trans. Amer. Philos. Soc., ser. 2 7:319. 1841. Haplopappus resinosus(Nutt.) A. Gray, Bot. Calif. 1:313. 1876. Chryso- thamnus resinosus (Nutt.) J.T. Howell, Fl. N.W. Amer. 303. 1900.
5. Ericameria watsonii (A. Gray) Nesom, comb. nov. BASIONYM: Hap- lopappus watsoni A. Gray, Proc. Amer. Acad. Arts 16:79. 1881. Mfacro- nema watsonu (A. Gray) E. Greene, Erythea 2:74. 1894.
D. Ericameria sect. Macronema (Nutt.) Nesom, comb. nov. BASIONYM: Macronema Nutt., Trans. Amer. Philos. Soc., ser. 2 7:322. 1841. Hap- lopappus sect. Macronema (Nutt.) A. Gray, Proc. Amer. Acad. Arts 6:542. 1865. TYPE SPECIES: Ericameria suffruticosa (Nutt.) Nesom.
1. Ericameria bloomeri (A. Gray) Macbr., Contr. Gray Herb. 56:36. 1918. Haplopappus bloomerit A. Gray, Proc. Amer. Acad. Arts 6:541. 1865. Chrysothamnus bloomert (A. Gray) E. Greene, Erythea 3:115. 1895.
2. Ericameria compacta (H.M. Hall) Nesom, comb. nov. BASIONYM: Haplopappus bloomeri A. Gray subsp. compactus H.M. Hall, Carnegie Inst. Washington, Publ. 389:199. 1928. Haplopappus compactus (H.M. Hall) L.C. Anderson, Great Basin Nat. 43:358. 1983.
3. Ericameria crispa (L.C. Anderson) Nesom, comb. nov. BASIONYM: Haplopappus crispus L.C. Anderson, Great Basin Nat. 43:359. 1983.
Nesom: Taxonomic summary of Ericameria 153
4a. Ericameria discoidea (Nutt.) Nesom, comb. nov. BASIONYM: Macro- nema discoidea Nutt., Trans. Amer. Philos. Soc., ser. 2 7:322. 1841. Haplopappus macronema (Nutt.) A. Gray, nom. nov., Proc. Amer. Acad.
Arts 6:542. 1865.
4b. Ericameria discoidea var. linearis (Rydb.) Nesom comb. nov. BA- SIONYM: Macronema linearis Rydb., Mem. New York Bot. Gard. 1:384. 1900. Haplopappus macronema (Nutt.) A. Gray var. linearis (Rydb.) Dorn, Vascular Plants of Wyoming 295. 1988.
5. Ericameria gilmanii (S.F. Blake) Nesom, comb. nov. BASIONYM: Haplopappus gilmanii S.F. Blake, Proc. Biol. Soc. Washington 52:97. 1939.
6. Ericameria greenei (A. Gray) Nesom, comb. nov. BASIONYM: Hap- lopappus greenet A. Gray, Proc. Amer. Acad. Arts 16:80. 1880. Afacrone- ma greenet (A. Gray) E. Greene, Erythea 2:73. 1894.
7. Ericameria ophitidis (J.T. Howell) Nesom, comb. nov. BASIONYM: Haplopappus bloomer: var. ophitidis J.T. Howell, Leaflets West. Bot. 6:85. 1950. Haplopappus ophitidis (J.T. Howell) Keck, Aliso 4:103. 1958.
8. Ericameria suffruticosa (Nutt.) Nesom, comb. nov. BASIONYM: Macronema suffruticosa Nutt., Trans. Amer. Philos. Soc., ser. 2 7:322. 1841. Haplopappus suffruticosus (Nutt.) A. Gray, Proc. Amer. Acad. Arts 6:542. 1865.
9. Ericameria zionis (L.C. Anderson) Nesom, comb. nov. BASIONYM: Haplopappus zionis L.C. Anderson, Great Basin Nat. 43:360. 1983.
Species Excluded
The following species will be treated as a separate genus (Nesom, et al. submitted).
Ericameria austroterana M.C. Johnston Ericameria diffusa Benth.
Ericameria parrasana S.F. Blake
Ericameria pseudobaccharis (S.F. Blake) Urbatsch Ericameria purpusti Brandegee
Ericamerza riskindit Turner & Langford
154 PHYTOLOGIA volume 68(2):144-155 February 1990
Ericameria triantha (S.F. Blake) Shinners
ACKNOWLEDGMENTS
I thank Dr. B.L. Turner and David Morgan for their review and comments on the manuscript.
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BOOKS RECEIVED
Annual Review of Entomology vol. 35. Thomas E. Mittler, ed. Annual Reviews, Inc., Palo Alto, CA, 1990. x. 641 pp. $38.00 (cloth).
Contribuciones de Eizi Matuda (1894-1978) al Conocimineto de la Flora de México, Cuadernos del Instituto de Biologia, U.N.A.M. vol. 1. Armando Butanda. Instituto de Biologia, U.N.A.M., Ciudad México, D.F., 1989. 67 pp. Price unknown.
Developmental Biology of Fern Gametophytes. V. Raghavan. Cam- bridge University Press, New York, NY, 1989. xiv. 361 pp. $80.00 (cloth).
Jerry Baker’s Flowering Garden. Jerry Baker. Collier Books, MacMillan Publishing Co., New York, NY, 1990. x. 229 pp. $7.95 (paper).
Morphology of Flowers and Inflorescences. F. Weberling. Cambridge University Press, New York, NY, 1989. xx. 405 pp. $110.00 (hardcover).
Native Shrubs and Woody Vines of the Southeast, Landscaping Uses and Identification. Leonard E. Foote & Samuel B. Jones, Jr. Timber Press, Inc., Portland, OR, 1989. 199 pp. $32.95 (hardcover).
Plant Stress from Air Pollution. Michael Treshow & Franklin K. Ander- son. John Wiley & Sons, Inc., Chichester, England, 1989. xi. 283 pp. $59.95 (hardcover).
Plants Under Stress, Society for Experimental Biology Seminar Series 39. Hamlyn G. Jones, T.J. Flowers & M.B. Jones, eds. Cambridge University Press, New York, NY, 1989. $65.00 (hardcover).
Textbook of Pollen Analysis, 4th ed. Knut Faegri & Johs. Iversen. John Wiley & Sons, Inc., Chichester, England, 1989. x. 328 pp. (price un- known).
The European Garden Flora III, Casuarinaceae to Aristolochiaceae. S.M. Walters, J.C.M. Alexander, A. Brady, C.D. Brickell, J. Cullen, P.S. Green, V.H. Heywood, V.A. Matthews, N.K.B. Robson, P.F. Yeo & S.G. Knees, eds. Cambridge University Press, New York, NY, 1989. (price unknown).
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Information for Authors
Articles from botanical systematics and ecology, including biographical sketches, critical reviews and summaries of literature will be considered for publication in PHYTOLOGIA. Manuscripts may be submitted either on computer diskette, or as typescript. Diskettes will be returned to authors after action has been taken on the manuscript. Diskettes may be 5.25 inches or 3.5 inches but must be written in DOS format or as flat ASCII files. Typescript manuscripts should be single spaced and will be read into the computer using a page scanner. The scanner will read standard typewriter fonts but will not read dot matrix print. Manuscripts submitted in dot matrix print cannot be accepted. Use underscore (not italics) for scientific names. Corrections made on typescript manuscripts must be complete and neat as the scanner will not read them otherwise. Language of manuscripts may be either English or Spanish. Figures will be reduced to fit within limits of text pages and therefore, should be submitted with an internal scale and have dimensions proportional to those for text pages. Legends for figures should be included in figures whenever possible. Each manuscript should have an abstract and key word list. Specimen citations should be consistent throughout the manuscript. Serial titles should be cited with abbreviations used in Botanico Periodicum Huntianum. References cited only as part of nomenclatural summaries should not appear in Literature Cited. © Nomenclatural work should include one paragraph per basionym — and must provide proper (as defined by the current /nternational Code of Botanical Nomenclature) citation of sources of epithets ~ and combinations. |
Authors should arrange for two workers in the appropriate field to review the manuscript before submission. Copies of reviews should ~ be forwarded to the editor with the manuscript. Manuscripts will not be published without review. 4
Cost of publication is currently $12.00 US per page for publica- — tion without reprints. Publication with 100 reprints is provided for — $16.50 US per page, 200 reprints for $20.00 US per page. Page ~ charges are due with manuscript and no paper will be published before payment is received in full. Reprints must be ordered and paid for in advance. Page charges will be determined on the basis ~ of a typescript page (single spaced, 10 points, blank line between — paragraphs) with all type inside a rectangle 143 mm (horizonal) by © 219 mm(vertical), not including running head and page number. Title page should include title, author(s) name(s) and address(es). Two blank lines should appear above and below section headings — (Abstract, Discussion, Literature Cited, etc.) in the manuscript. No — extra charge is made for line drawings provided they conform to- limitations of size and proportion for normal text. Halftones require an extra charge of $5.00 US per page.