" PHYTOLOGIA An international journal to expedite plant systematic, phytogeographical and ecological publication Vol. 81 July 1996 No. 1 CONTENTS TURNER, B.L., A new species of. Geranium (Geraniaceae) from Oaxaca, MPN a eaily COMO ce inane Der eles. SPaviy pee tT Mitagae OC tye SE MCR Rae Amok grr Sante te pe! DENHAM, R.A. & B.L. TURNER, A new species of Tetraneuris (Asteraceae, Helenieae) from the late Tertiary Verde Formation of central Anizona......... hie TURNER, B.L., Synoptical study of the Acacia angustissima es se COMBE R Se aes acs vv oak eaask a vig Phase Pade sh Pan yah be oe eLeeawsenie Lus ea TURNER, B.L., Synopsis of section Axillaris of Salvia (Lamiaceae).......:.. i 3 TURNER, B.L., Russelia mananilana (Scrophulanaceae), a new. species Ligh Jalisco, Weide so ie Se eR ae Sp HENSOLD, N., Paepalanthus subg. Xeractis (Enocaulaceae): Notes ad ee nomenclatural NAN BCS 5055 ¢ is torts hr tears scope tert a. Yerial cs tnstahiaivedais 24 MACROBERTS, M.H.. & B.H: MACROBERTS, Lorgleaf pine (Pinus palustris: Mill. ¥-erowth 10) DOSS nS Ooch red cl ks os. Sig ees be 28 MACROBERTS, B.R. & M.H. MACROBERTS, The floristics of calcareous praines on the Kisatchie National Forest, Louisiana. ................2....0... 35 a SMITH, E.B., Comments on the “Coreocarpus arizonicus-C. sonoranus | (Asteraceae), Heliantheae) complex”. «22... 0.00... cece cece eee eee cc cn ees 44 ROSALES-CARRILLO, O. & Y. HERRERA-ARRIETA, Epidermis en laminas foliares del género Bouteloua Lagasca, (Poaceae: Chlioridoideae) de México Index to authors, volume 80. ..... 000.00. -.g. pg. gem om. A. PMR... 65 Index to reviewers, volume 80. ......:.... \.. [ Fee ARY ine woken 66 Publiction dates. VOLUME Bl, pees reat eee oC a ee kas 67 Prytoloma Memos at available: ios oa es ceeds biti deo sno aes 68 MAY 5 ~ 1997 NEW YORK SUTANICAL GARDEN Published by Michael J. Warnock 185 Westridge Drive Huntsville, Texas 77340 U.S.A. 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Westridge — Drive, Huntsville, TX 77340-8916, gee Phytologia (July 1996) 81(1):1-4 A NEW SPECIES OF GERANIUM (GERANIACEAE) FROM OAXACA, MEXICO B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A new species of Geranium, G. miahuatlanum B.L. Turner, is described and illustrated from Mpio. Miahuatlan, Oaxaca, México. It belongs to the senes Repentia where it relates closely to G. repens, a species of Central America, occurring from Guatemala to Panama. KEY WORDS: Geraniaceae, Geranium, México, Oaxaca Geranium is a difficult genus, especially in México where numerous, closely related, taxa are recognized. Moore (1943) provided the last definitive treatment of the genus for México, in which 40 or more species were recognized, distributing these among twelve weakly defined series. The present novelty belongs to his series Repentia where it relates most closely to G. repens H. Moore. GERANIUM MIAHUATLANUM B.L. Turner, spec. nov. Figure 1. TYPE: MEXICO. Oaxaca: Mpio. Miahuatlén, San Agustin, 2735 m, swamp in pine forest, forming colonies, 5 Aug 1996, Hinton et al. 26744 (HOLOTYPE: TEX!). Similis G. repenti H. Moore sed foliis majonbus, lobis mediis plerumque 2.5-3.5 cm longis (vice 1.8-2.3 cm longis); pedicellis longioribus, plerumque 2.0-3.5 cm longis (vice plerumque 0.5-2.5 cm longis); et calycibus majoribus, tubis 6-7 mm longis (vice 5-6 mm longis). Repent herbs 0.3-0.6 m high. Stems moderately pubescent with spreading, glandular or eglandular hairs to 1.2 mm long; internodes elongate 2-5 times longer than the midstem leaves. Midstem leaves deeply 3(-5)-cleft, the midlobes 2.5-3.5 cm long, somewhat longer than the lateral lobes. Peduncles mostly 3-6 cm long, 2- flowered, glandular-pubescent. Sepals 7-9 mm long, the bodies 6-8 mm long, glandular-pubescent, 3-nervate. Petals 12-13 mm long, pink to dark pink, glabrous except for the ciliate claw. Capsules ca. 2.5 cm long, the beaks ca. 3 mm long. to PHYTOLOGIA July 1996 volume 81(1):1-4 ADDITIONAL SPECIMENS EXAMINED: MEXICO. = Oaxaca: Mpio. Miahuatldn, Neverias, 2650 m, 1 Jun 1996, Hinton et al. 26552 (TEX); San Agustin, 2750 m, 5 Aug 1996, Hinton et al. 26759 (TEX). The elongate internodes, larger leaves with more linear-lanceolate lobes, longer pedicels and more deeply pink petals readily distinguish this species from its most closely related cohort, Geranium repens. The latter, so far as known, is confined to the montane regions of Central America from Guatemala to Panamé, although Moore (1943) cited a single specimen from Guerrero, México, which I take to be a misidentification of G. hintonii H. Moore (discussed below). The series Repentia, as treated by Moore (1943) contains three species: a widespread Geranium repens; G. clarum Small, from Oaxaca, known only from type material; and G. hintonii, from Guerrero, known only from type matenal. The following key will distinguish among the four taxa which make up the series Repentia, as recognized here. KEY TO MEXICAN TAXA OF SERIES REPENTIA 1. Pedicels eglandular-pilose; petals white to pale pink. 2. Pedicels 5-10 mm long; northcentral Oaxaca. ...........-....0seeeeee ee G. clarum 2. Pedicels 12-30 mm long; central Guerrero.................::::e ee G. hintonti 1. Pedicels glandular-pilose; petals pink to dark pink. 3. Middle lobe of mid-stem leaves 2.5-3.6 cm long; pedicels mostly 25-35 mm long; body of calyx 6-8 mm long; Oaxaca..............:-.2+ G. miahuatlanum 3. Middle lobe of mid-stem leaves 1.8-2.2 cm long; pedicels mostly 20-25 mm long; body of calyx 5-6 mm long; Guatemala to Costa Rica........... G. repens As noted in the above, Moore included in his concept of Geranium repens a questionable plant from Guerrero, México, “Hinton 14579’ which he commented upon, as follows: “The northern representative, Hinton 14579, from Guerrero, has villous pubescence and large petals but is so similar in other respects that I am placing it here at present.” Actually the collection number concerned, in my opinion, should be Hinton 14759, as shown by two collections at LL!, TEX! and as can be vouched for in Hinton & Rzedowski (1975). Regardless, Hinton 14759, in all its characters, clearly belongs to Geranium hintonii. Indeed, the plant itself was collected fairly near the type locality of the latter. ACKNOWLEDGMENTS I am grateful to Gayle Turner for the Latin diagnosis, and to her and Ted Delevoryas for reviewing the manuscript. Maria Thompson provided the illustration. Turner: New Geranium from Oaxaca Lem Figure 1. Geranium miahuatlanum (from holotype). 4 PHY TOLGIGLTA July 1996 volume 81(1): 1-4 LITERATURE CITED Hinton, J. & J. Rzedowski. 1975. George B. Hinton, Explorador Botdnico en el Sudoeste de México. An. La Escuela Nac. Cien. Biol. 21:1-114. Moore, H.E., Jr. 1943. A revision of the genus Geranium in Mexico and Central America. Contr. Gray Herb. 146:1-108. Phytologia (July 1996) 81(1):5S-9 A NEW SPECIES OF TETRANEURIS (ASTERACEAE, HELENIEAE) FROM THE LATE TERTIARY VERDE FORMATION OF CENTRAL ARIZONA Robert A. Denham! & B. L. Tuner ‘Current address: 3609 W. Jasmine Rd., Las Cruces, New Mexico 88005 U.S.A. *Department of Botany, The University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A new species, Tetraneuris verdiensis R.A. Denham & B.L. Turner, is descnbed from Yavapai County, Anzona where it is restricted to lacustrine marl in the Verde Formation. It is related to the Tetraneuris “scaposa -- acaulis complex,” and can be readily distinguished from the closely sympatric T. scaposa (Hymenoxys acaulis var. arizonica) by its dwarf habit, relatively short broad leaves, long-pilose vestiture, and rayless heads. KEY WORDS: Asteraceae, Helenieae, Tetraneuris, Anzona Preparation of a systematic study of the genus Tefraneuris (Bierner & Tumer 1997) by the junior author has occasioned the present paper. This new species was brought to his attention by the senior author, who first became aware of its existence during the winter of 1993-1994 while engaged in flonistic studies of localized substrates within the lacustrine limestone Verde Formation in northeastern Yavapai County, Anzona. Jean Searle of the Amzona Native Plant Society, while accompanying the senior author on a field trip to the type location, first pointed out the uniqueness of these populations as compared to Tetraneuris elsewhere in Anzona. TETRANEURIS VERDIENSIS R.A. Denham & B.L. Turner, spec. nov. TYPE: U.S.A. Anzona: Yavapai Co., 5 mi. E of Camp Verde, 3300 ft., occurring on marl with gypsum crystals at the surface, 14 May 1995, Denham, Fobes, & Searle 1840 (HOLOTYPE: TEX). Similis Tetraneuri scaposae (DC.) Greene sed planta nana et eradiata est, 4-7 cm alta, indumentum candudum -- pilosum habens, pilis 3-7 mm longus. Dwarf scapose perennial 4-7 cm high, the stems ansing from a branched woody caudex. Leaves relatively thick, all basal, 1.5-2.5 cm long, 3-5 mm wide; blades ovate to narrowly ovate, moderately but deeply glandular-punctate, entire, markedly 5 6 PHy TOLOGTIA July 1996 volume 81(1):5-9 white-pilose throughout with hairs 3-6 mm long, the apices acute to obtuse. Scapes 4- 6 cm long, ebracteate, pilose with upwardly appressed and widely spreading hairs 1-4 mm long. Heads single, hemispheric. Involucres 5-6 mm high, 3-4 mm across. Ray florets absent. Disk florets ca. 40 (est.); corollas yellow, ca. 3.5 mm long, the tube ca. 0.5 mm long, the lobes 5, densely pubescent. Anthers yellow with ovate appendages. Style branches truncate, apically hispidulous. Achenes (immature) ca. 3 mm long, densely pubescent; pappus of ca. 8 lanceolate scales 2-3 mm long. Additional collections: Denham, Fobes, & Searle 1835, 1836, 1837, 1838, and 1839, all from the same location as the type, from either the same population as the type or from an adjacent population on a nearby hilltop. GEOLOGY AND FLORISTICS OF THE TY PE LOCATION Tetraneuris verdiensis can best be understood with a perspective on the geology and floristic patterns of the type location and surrounding areas within the Verde Formation. During the late Tertiary Period, the Verde Formation and other lacustrine deposits formed in a series of basins across the sub-Mogollon region of Anzona. These basins are home to a number of endemics which are often restricted to particular substrates. In addition to these endemics, disjunct occurrences of several other species are found in these lacustrine deposits (Anderson 1996). The Verde Formation in northeastern Yavapai County was formed pnmanily through deposition and precipitation within a shallow lake bed created by down- dropping along the Verde Fault and subsequent blockage of the drainage outlet by volcanic and/or tectonic activity. The upper part of the formation, exposed at the northwestern end of the Verde Valley near Cottonwood, Anzona, is comprised of narrow interbedded layers of limestones, mudstones, and marls. The lower part of the formation, exposed at the southeastern end of the valley near Camp Verde, Arizona, is comprised mostly of more massive limestones formed in the deepest parts of the lake and evaponites, such as salt deposits and gypsum. Near the lower end of the valley are volcanoclastic deposits adjacent to and of approximately the same age as the lacustrine deposits. Major changes in the floristic communities occur along with changes in substrate within the Verde Formation. The xeric hillside habitats on the interbedded layers near the upper end of the valley are dominated by Canotia holacantha and Juniperus coahuilensis (Martinez) Gaussen ex R.P. Adams. On the massive limestones in the lower valley, the flora includes some species more typical of higher elevations, such as Juniperus osteosperma (Torrey) Little, Purshia stansburiana (Torrey) Henrickson, and Ipomopsis aggregata (Pursh) V. Grant. The adjacent volcanic tuff supports a community with a Sonoran component, including Agave chrysantha Peebles, Opuntia acanthacarpa Engelm. & Bigelow var. thornberi (Thornber & Bonker) L. Benson, and Acacia constricta Benth. Several taxa endemic to the late Tertiary sub-Mogollon lacustrine basins are found to occur along particular soil honzons. In the Verde Valley, Purshia subintegra New Tetraneuris from Arizona Denham & Turner: lotype. The Tetraneuris verdiensis R.A. Denham & B.L. Turner, from ho enlarged circle at lower right depicts several folded leaves. Figure 1. 8 PHYTOLOGIA July 1996 volume 81(1):5-9 (Kearney) Henrickson only occurs near the upper end of the Verde Formation and then only where there are clastic elements derived from the Mesozoic sandstones of the Supai Group, and Eriogonum apachense is restricted to a horizon at ca. 3500' in the block-forming limestones of the lower valley. Many other species with disjunct occurrences in the lacustrine Verde Formation also follow a similar pattern of being restricted to specific soil honzons. For example, Quercus havardii Rydb. var. tuckeri Welsh from southeastern Utah is disjunct from its nearest population by almost 170 miles, and is found in central Anzona only at locations where a particular powdery clay-like calcium carbonate soil is exposed at ca. 3400' in the vicinity of Dead Horse Ranch State Park. In the lowest, southeastern-most end of the lacustrine deposits, several narrow bands of different soils running northeast-southwest are exposed. Within three miles, one can go from limestones, across bands of gypsum and marl, and continue onto volcanic tuff, each of these substrates accommodating their own floristic community. In this area, the type population of Tetraneuris verdiensis occurs on one of a series of low chalky flat-topped hills composed of marl with gypsum crystals at the surface. Additional populations of Tetraneuris verdiensis occur on the tops of the adjacent small hills within this stratigraphic unit. The type location supports a dwarf sub-shrub community dominated by Eriogonum ericifolium Torrey & A. Gray var. ericifolium, an edaphic endemic known only from lacustrine deposits within the Verde Formation. Co-dominant is Salvia dorrii (Kell.) Abrams subsp. mearnsii, endemic to the Verde Formation and adjacent sandstones. Overall, the distribution of Tetraneuris verdiensis is consistent with the pattern of edaphic endemism found within the Verde Formation and other late Tertiary lacustrine deposits in central Anzona. THE TAXONOMY OF TETRANEURIS VERDIENSIS The genus Tetraneuris is known from the Great Basin, the Rocky Mountains, and the Great Plains. It reaches its southwestern limit in north-central México. Tetraneuris scaposa Greene (including Hymenoxys acaulis (Pursh) Parker var. arizonica [Greene] Parker) is common in the northern parts of Anzona. This taxon also has a disjunct range to the south in the lacustrine limestone Verde Formation and adjacent Mesozoic sandstones in Yavapai County (Anderson 1996 ibid.). In the Verde Valley, T. scaposa has been collected by the senior author at Cottonwood, Anzona near the upper end of the valley. Anderson (1996 and pers. comm.) has collected this same taxon along Middle Verde Rd., west of Camp Verde, Arizona. This latter site is approximately seven miles from the type location of T. verdiensis. Although both of these species occur in the Verde Valley, there is no evidence of intergradation, no intermediate forms, and no individuals exhibiting a recombination of characteristics between these two species. _ _One important characteristic of Tetraneuris verdiensis is its discoid heads. Discoid individuals are known elsewhere in Tetraneuris, as isolated individuals within populations of 7. acaulis Greene (Hymenoxys acaulis var. acaulis) in Wyoming. Denham & Turner: New Tetraneuris from Arizona 9 These individuals, which have formed the basis for 7. eradiata A. Nelson, differ from their neighbors only in their discoid condition, and are currently considered to be aberrant forms of 7. acaulis. The situation in the Verde Valley is radically different. Here 7. scaposa and T. verdiensis can be distinguished by a suite of characteristics which are always consistent at the population level. In addition to its discoid heads, T. verdiensis differs from T. scaposa in its dwarf habit, relatively short broad leaves, and long pilose vestiture. Other than 7. verdiensis, there has been no report in Tefraneuris of entire populations, or series of populations, which are wholly discoid. CONCLUSION In Tetraneuris and in the closely related Hymenoxys, a syndrome of charactenstics, some of these subtle, separate the various species. The degree of morphological distinction of Tetraneuris verdiensis is consistent with recognition at the species level within both of these genera. In short, Tetraneuris verdiensis is a relatively well-marked localized edaphic endemic of central Arizona. ACKNOWLEDGMENTS Gayle Turner provided the Latin diagnosis, and we are grateful to Mark Bierner and Ted Delevoryas for reviewing the manuscript. We would also like to thank John _L. Anderson for additional information on the distnbution of Tetraneuris scaposa (Hymenoxys acaulis var. arizonica) in the Verde Valley, and Elizabeth Mathews, Northern Anzona Zone Geologist, United States Forest Service, for clarification of the finer points of the geology of the Verde Valley. LITERATURE CITED Anderson, J.L. 1996. Floristic Patterns on Late Tertiary Lacustrine Deposits in the Arizona Sonoran Desert. Madrofio 43:225-272. Biermner, M. & B.L. Turner. 1997. Systematic overview of the genus Tetraneuris (Asteraceae: Helenieae). in prep. Phytologia (July 1996) 81(1):10-15. SYNOPTICAL STUDY OF THE ACACIA ANGUSTISSIMA (MIMOSACEAE) COMPLEX B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A taxonomic study of the widespread North Amenican species, Acacia angustissima, is rendered. It is recognized as having six intergrading morphogeographical infraspecific categories, as follows: 1.) var. angustissima, a shrub or small tree occurring in tropical and subtropical mesic habitats along both sides of México from the states of Jalisco and Nuevo Leon southwards to Panamé; 2.) var. hirta, a rhizomatous perennial herb with simple stems occurring in the southeastern and south-central U.S.A.; 3.) var. suffruticosa, a suffruticose herb or shrub occurring mostly in the Sonoran desert regions of northeastern México; 4.) var. chisosiana, a divaricately branched subtortuose shrublet or shrub of north-central México and closely adjacent U.S.A. (New Mexico and trans-Pecos Texas); 5.) var. leucothrix (Standl.) B.L. Turner, comb. & var. nov., a divaricately branched shrublet or shrub of the Tamaulipan scrublands of northeastern México and closely adjacent U.S.A.; and 6.) var. oaxacana B.L. Turner, var. nov., a shrub or shrublet of the xeric scrublands of southern Pueblo and northern Oaxaca, México. A key to the varieties and a map showing their distnbution is provided. KEY WORDS: Mimosaceae, Acacia, U.S.A., México Acacia angustissima (Mill.) Kuntze is a widespread common species of North Amenica. It has been the subject of three relatively detailed studies (Wiggins 1942; Isely 1969; McVaugh 1987). Isely especially clarified relationships in the complex in northern México and the U.S.A. with his recognition of six intergrading varieties: angustissima, hirta (Nutt.) B.L. Rob., texensis (Torrey & A. Gray) Isely, chisosiana Isely, suffrutescens (Rose) Isely, and shrevei (Bntt. & Rose) Isely. Of these, I combine var. fexensis with var. hirta, and var. shrevei with var. suffruticosa. | have also recognized two additional infraspecific taxa, var. leucothrix and var. oaxacana. A key and map (Figure 1) to these various taxa follows, along with justification for the treatment, including partial synonymies where pertinent. More detailed information re Tumer: Synopsis of Acacia angustissima complex 11 nomenclature and typification can be found in the papers of Wiggins (1942), Isely (1969), and McVaugh (1987). Except as otherwise noted, the map (Figure 1) is based upon the several hundred specimens of the complex on file at LL, TEX. All of these have been appropnately annotated, including those of an intergradant nature. KEY TO VARIETIES OF ACACIA ANGUSTISSIMA 1. Leaves with mostly 10-20 pairs of pinnae; plants of south-central and southeastern U.S.A. or tropical and subtropical México to Central Amenica. .................... (2) 2. Suffruticose herbs mostly 0.3-1.0 m high; south-central and southeastern LE Ae ta saeco Getreice «asin: Solan snap ata cates cag SeRaeh a See eer aeraeee var. hirta 2. Shrublets, shrubs, or small trees mostly 2-7 m high. .......... var. angustissima 1. Leaves with mostly 2-9-10) pairs of pinnae; plants of more xeric montane or subdesert habitats, southwestern U.S.A., Sonoran and Chihuahuan deserts, Tamaulipan biotic province, and Puebla-Oaxacan shrublands..................066 (3) 3. Pinnae mostly 1-2 cm long; much-branched subtortuous shrublets of Chihuahuan Desert and Tamaulipan shrublands.....................0.0.c0eeeeeee (4) 4. Older stems markedly striate with white to tan grooves; Chihuahuan Desert regions or north-central México, and closely adjacent New Mexico and MIGXAS, S50. ao csc veces cose pasens an cavepeas teeacaencsteecwesGeeees var. chisosiana 4. Older stems only weakly strate if at all; Tamaulipan biome scrublands of northeasternmost México (Tamaulipas and Nuevo Leén) and closely ARACETE LOXAS oe Geeta ns's eeainan de cena sc naeeebisen ans os abapesnts var. leucothrix 3. Pinnae of well-developed leaves mostly 2-3 cm long; mostly straight-stemmed or weakly subtortuose suffruticose herbs or shrublets of the Sonoran Desert and Pacific slopes, or xeric montane habitats of southern Puebla and northern Oaxaca’, cn sienr de Rewer ee oO OO ee a ee ne ore (5S) 5. Pinnae mostly 2-4(-8) pairs; shrublets with subtortuous stems; southern Pueblaiandinorthem @axacas=+.-0-.-peeee se eneee ee ere ee eee var. oaxacana 5. Pinnae mostly (4-)6-9(-10) pairs; suffruticose herbs to shrubs with mostly nontortuose stems; Sonoran Desert regions and Pacific slopes from Sonora to Nayant and northern Jalisco. ...............cceceeeeeee ens var. suffrutescens ACACIA ANGUSTISSIMA (Mill.) Kuntze var. AVNGUSTISSIMA Acacia angustissima (Mill.) Kuntze, Rev. Gen. Pl. 3:47. 1896. Mimosa angustissima Mill., Gard. Dict. ed. 8. Mimosa no. 19. 1768. TYPE: MEXICO. Veracruz: 1731, Houston s.n. (BM), according to Mc Vaugh (1987) This is the most widespread and frequently encountered vanety of the Acacia angustissima complex, occurring in tropical and subtropical habitats from northeastern Nuevo Le6én, México and extending down the Gulf Coastal slopes to Panama; on the Pacific slopes it occurs from southern Durango and probably closely adjacent Sinaloa to Panama (Figure 1). The taxon is readily recognized by its large habit (2-8 m), 12 PHYROELOGTA July 1996 volume 81(1):10-15 elongate nontortuose stems, and large leaves with numerous pinnae (mostly 10-20 pairs). Wiggins (1942) provided a partial list of synonyms for the taxon (including Acacia angustissima subsp. smithii [Britt. & Rose] Wiggins, which I cannot recognize), as did McVaugh (1987). Additional synonyms are likely to be disinterred from among the numerous names proposed for the North American elements of Acacia. The type of A. angustissima was obtained from Veracruz, México by Houston in 1731 and is discussed in more detail by Wiggins (1942). ACACIA ANGUSTISSIMA (Mill.) Kuntze var. CHISOSIANA Isely Acacia angustissima (Mill.) Kuntze var. chisosiana |sely, Sida 3:370. 1969. Isely, although not having examined living plants, recognized the distinctiveness of this taxon. Turner (1959) erroneously included most such material in his concept of Acacia texensis Torrey & A. Gray. The latter name is typified by material from near New Braunfels, Texas, and is nothing more than forms of var. hirta, having somewhat fewer pinnae. The var. chisosiana is largely confined to the Chihuahuan Desert regions of trans- Pecos Texas, New Mexico, and north-central México (Figure 1). It appears to intergrade but slightly into var. /urta in the eastern portion of its range and perhaps into var. suffrutescens in the western parts of its range. Indeed, it is possible that future workers might consider it specifically distinct; if so, this would perhaps necessitate the elevation of varieties /eucothrix and oaxacana, this tnad being superficially similar among themselves, but unlikely to represent a monophyletic element, to judge from their distributions. ACACIA ANGUSTISSIMA (Mill.) Kuntze var. HIRTA (Nutt.) B.L. Rob. Acacia hirta Nutt., in Torr. & Gray, Fl. N. Amer. 1:404. 1840. Acacia angustissima (Mill.) Kuntze var. hirta (Nutt.) B.L. Rob., Rhodora 10:33. 1908. Type collected by Nuttall in eastern Oklahoma or closely adjacent Arkansas. Acacia texensis Torr. & Gray, Fl. N. Amer. 1:404. 1840. Acacia angustissima (Mill.) Kuntze var. fexensis (Torr. & Gray) Isely, Sida 3:372. 1969. Type collected by Lindheimer in central Texas, vicinity of New Braunfels, Comal Co. Isely (1969) recognized Acacia texensis as varietally distinct, distinguishing it from var. hirta by its purportedly smaller leaves with fewer pinnae (mostly 4-6 pairs vs. 9- 15 pairs), ignoring intermediates between these. I view Isely’s var. lexensis to be but sporadic leaf forms of the widespread var. /iirta, the smaller leaves usually occurring on secondary shoots which arise from cutover or browsed plants, although some such plants must represent intermediates between var. chisosiana and var. hirta along their regions of contact. a 200 Km A var. ANGUSTISSIMA @ var. CHISOSIANA O var. HIRTA oO Pp @ var. LEUCOTHRIX var. OAXACANA var. SUFFRUTICOSA Synopsis of Acacia anguslissima complex 13 Wineney SP Gadi TSS a SE ae me alte 3 BET LIAL it mall rae : : {hfe oe Bi) os wo e am ye) ow —To] “La tH - ene QPi9. oy er Figure 1. Distnbution of the infraspecific categones of Acacia anguslissima. All of these apparently intergrade to some extent in areas of close proximity, but only the half-closed circles documented this for var. chisosiana and var. hirta is shown on the map. Arrows show extension of the taxa to regions outside of the bounds of the map area. The map is based largely upon specimens at LL, TEX, except for most of the open circles in Oklahoma and Nebraska, these having been extracted from the map portrayed in Barkley (1977). 14 PHY TOLOIGITA July 1996 volume 81(1):10-15 ACACIA ANGUSTISSIMA (Mill.) Kuntze var. LEUCOTHRIX (Standl.) B.L. Turner, stat. & comb. nov. BASIONYM: Acacia leucothrix Standl., Contr. U.S. Natl. Herb. 20:185. 1919. The type was collected in easternmost San Luis Potosf, México (at San Dieguito) in 1904 by Palmer. (Paratypes examined: Pringle 9717 (3 sheets, LL, TEX)). This taxon is well-represented at LL, TEX. Dr. M.C. Johnston was perhaps the ' first to recognize its affinity, at least by annotations on several sheets at LL, TEX. Standley in his protologue compared his new taxon with Acacia cuspidata Schlecht., but the latter presumably does not belong to the A. angustissima complex if we are to believe that the leaf petiole is glandular, as given in its type description. ACACIA ANGUSTISSIMA (Mill.) Kuntze var. OAXACANA B.L. Turner, var. nov. TYPE: MEXICO. Oaxaca: 5km al SE de Cuicatlan, por la desviacion a San Pedro Ocotipac, Selva Baja Caducifolia, ca. 760 m, 27 Aug 1980, F.G. Madrano, et al. F-1568 (HOLOTY PE: TEX!). Similis A. angustissimae (Mill.) Kuntze var. leucothrix (Standl.) B.L. Turner sed habens folia majora cum pinnis longioribus (2-3 cm vice 1-2 cm) et caules minus tortos. ADDITIONAL SPECIMENS EXAMINED (9 sheets): MEXICO. Puebla: Salinas T. & Dorando R. F-3097 (TEX); Sousa 9390 (TEX); Tenorio L. 8014 (TEX), Tenorio L. 14158 (TEX). Oaxaca: Magallanes 53, 196 (TEX); Salinas T. 4635, 4846, 4861 (TEX). This taxon is superficially similar to var. leucothrix but has mostly larger leaves with longer pinnae (2-3 cm long vs. 1-2 cm long) and less tortuose stems. The collections by Magallanes (cited above) differ from most of the other collections in having mostly 4-8 pairs of pinnae, otherwise the plants are scarcely distinguishable. ACACIA ANGUSTISSIMA (Mill.) Kuntze var. SUFFRUTESCENS (Rose) Isely Acacia suffrutescens Rose, Contr. U.S. Natl. Herb. 12:409. 1909. Acacia angustissima (Mill.) Kuntze var. suffrutescens (Rose) Isely, Sida 3:372. 1969. The type was collected in Santa Cruz Valley, near Tucson, Arizona, U.S.A. Acacia lemmonii Rose, Contr. U.S. Natl. Herb. 12:409. 1909. — Acacia angustissima (Mill.) Kuntze subsp. lemmonii (Rose) Wiggins, Contr. Dudley Herb. 3:230. 1842. The type was collected by Lemmon in the Huachuca Mts., Cochise Co., Arizona, U.S.A. Acaciella shrevei Britt. & Rose, N. Amer. Fl. 23:105. 1928. Acacia angustissima (Mill.) Kuntze var. shrevei (Britt. & Rose) Isely, Sida 3:371. 1969. The type was collected by Shreve in the Huachuca Mts., Cochise Co., Arizona, U.S.A. As noted in the above synonymy, Isely (1969) recognized a var. shrevei from among this complex, largely distinguished by its purportedly shrubby habit and more Tumer: Synopsis of Acacia angustissima complex 1S venose leaflets. [I cannot see that such habitat forms might be meaningfully segregated, nor does leaflet venation serve to mark the Huachuca Mt. specimens as distinct, there being much sporadic variation of this character, especially in México. Indeed, McVaugh (1987) calls attention to similar venose forms in western México; he called such matenal var. texensis (Torr. & Gray) Isely. It is likely, however, that the plants concemed are southern elements of my concept of var. suffrutescens, the latter presumably intergrading into var. angustissima in this region. It should also be noted here that Acaciella painteri Britt. & Rose, discussed by McVaugh (1987, p. 125) presumably belongs to the var. suffruticosa complex. The limits of the latter taxon in Sinaloa, Durango, Zacatecas, Nayarit, and Jalisco is in much need of additional study, especially populational investigations. ACKNOWLEDGMENTS I am grateful to Gayle Tumer for the Latin diagnosis, and to her and Ted Delevoryas for reviewing early drafts of the manuscript. LITERATURE CITED Barkley, T.M. [ed.]. 1977. Atlas of the Flora of the Great Plains. lowa St. Univ. Press, Ames, Iowa. Isely, D. 1969. Legumes of the United States: I. Native Acacia. Sida 3:365-386. McVaugh, R. 1987. Leguminosae. Fl. Novo-Galiciana 5:1-786. University of Michigan Press, Ann Arbor, Michigan. Turner, B.L. 1979. The Legumes of Texas. Univ. of Texas Press, Austin, Texas. Wiggins, I.L. 1942. Acacia angustissima (Mill.) Ktze.: and its near relatives. Contr. Dudley Herb. 3:227-239. Phytologia (July 1996) 81(1):16-21 SYNOPSIS OF SECTION AX/ILLARIS OF SALVIA (LAMIACEAE) B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT The wholly Mexican sect. Axillaris Epling of the genus Salvia is revised. It is treated as having a single species, S$. azullaris Benth., with three morphogeographical infraspecific units: var. axillaris of southern Puebla and closely adjacent Oaxaca; var. hidalgoana B.L. Turner, var. nov., of southern Hidalgo; and var. potosina B.L. Turner, var. nov., of Durango, Zacatecas, Aguascalientes, San Luis Potosf, Guanajuato, Querétaro, and southwestem Hidalgo. An illustration of the species and a key to the varieties is provided, along with a map showing their distributions. KEY WORDS: Salvia, sect. Axillaris, Lamiaceae, México, systematics Attempts to identify various species of Salvia from México has led to the detailed examination of S. axillaris Benth., a very distinct species which was treated by Epling (1937) as belonging to the monotypic section Axillaris. Results of this investigation follow. SECTION AXILLARIS Epling (1937) Subsection Axilliflorae Benth., Lab. Gen. et Sp. 270. 1833. SALVIA AXILLARIS Moc. & Sesse ex Benth., Lab. Gen. et. Sp. 270. 1833. My inclusive description of this species is about the same as that provided by Epling (1937) and no redescription is needed here. I have, however, recognized three morphogeographical varietal elements within the taxon. Some workers might prefer to treat these as distinct species, especially since they occupy different nonconuguous ecogeographical settings and scarcely can be said to intergrade. There can be no Turner: Synopsis of Salvia axillaris 17 question, however, that these several taxa might not be more closely related one to the other than to yet other species. KEY TO VARIETIES 1. Filaments of stamens pubescent with coarse hairs; stem vestiture of mostly SPlANUUlar SPLCAGING MANES. rates cin os aereclss a cos udebeoatees sacanes aster var. potosina 1. Filaments of stamens glabrous (rarely a few basal hairs); stem vestiture of long glandular hairs, or short eglandular, mostly down-curved or arcuate hairs. 2. Vestiture at midstem mostly composed of spreading glandular trichomes 0.3- 0.5 mm high; midstem leaves weakly nervate, mostly 5-7 mm long, 2-3 times asloneas Wide Hidalgo: . 3.5...... ‘9 :sepsad * (00101 aa (6uauds) tAuued cal Fs] ae ofa -[92 * y Bouteloua epidermal anatom -A.: -C. & Herrera Rosales SEO costal intercostal zona Rosales-C. & Herrera-A.: Bouteloua epidermal anatomy 59 yeysoduaqut euoz ————! ye sod eudZz— | ~ < Fig. 4. B. hirsuta Lagasca var. hirsuta (De la Cerda de corcho; cie, células interestomatales; cl, células largas; cs, Cuer- pos de sf{lice; e, estomas; mi, micropelos. 400X. PHYTOLOGIA Bouteloua trifida Thurber in BL Ei, Cm ‘ eae at * Chae, » : a es . . k 4a am i Ae Ee . vas volume 81(1):46-64 Rosales-C. & Herrera-A.: Bouteloua epidermal anatomy DEXDROGRAMA . INDICE DE BUCLIDIAN MORMALIZADO avs B. disticha | : 27 3. parryi var. parryi = 39 B. uniflora var. uniflora 2 B. americana 14.8. distans | 6 B. parryi var. gentryi 38 B. uniflora var. coaluilensis 2B. curtipendula var. teuis (— B. trifida SB. media. 3B. annua | 16 B. elata | 28 B. pedicellata B. warnockii aan 24 as 4B. aristidoides var. aristidoides 5 B. barbata var. barbata fone 6 B. barbata var. rothrockii en B. eriopoda 38 B. radicosa 17 B. eludens L= B. quiriegoensis B. williansii 7 B. barbata var. scnorae | (a B. eriostachya 31 B. reflexa 8 B. breviseta | foee B. gracilis 32 B. repens 1 B. alamosana | 11 B. curtipendula var. caespitosa | oe B. johnstonii 35 B. Simplex 18 B. chondrosioides | = B. hirsuta var. hirsuta 34 B. scorpioides 9 B. chasei | 1 B. hirsuta var. glandulosa 33 B. rigidiseta ae B. curtipendula var. curtipendula es B. karwinskii 36 B. triaena 62 PHY TOLOGIA July 1996 volume 81(1):46-64 Las especies y variedades revisadas tienen caracteristicas comunes como son: presentar células alargadas paralelas horizontalmente, sinuosas; células cortas en pares (la mayorfa); estomas en forma de domo bajo, rectangulares y en hileras definidas en todas las zonas intercostales; micropelos bicelulares, cuerpos de sflice en forma de silla de montar formando una doble equis. Es conveniente explicar que las aseveraciones anteriores respecto a los ganchos son de acuerdo a la parte de donde se hizo el corte, ya que pueden existir variaciones en la misma hoja 0 en hojas diferentes de la misma planta, también dependiendo de la maduréz de éstas. DISCUSION Para llevar a cabo el andlisis fenético se tomaron once caracteres (Cuadro 2), de las 34 especies y doce variedades mexicanas del género Bouteloua. Con ellos se generé una matriz (Cuadro 3), con la cual se corrio un dendrograma, en el se observa como se van uniendo las especies por medio de nudos de acuerdo a las semejanzas que hay entre ellas y separando a las especies que comparten menos caracteristicas. [En el {indice de similitud Euclidiana Normalizada se agrupan las especies en tres grupos bien definidos. En el grupo I se incluyen las siguientes especies: B. uniflora var. coahuilensis, B. parryi var. gentryi, B. distans, B. americana, B. warnockii, B. trifida, B. media, B. curtipendula var. tenuis, B. parryi var. parryi, B. uniflora var. uniflora, B. disticha B. pedicellata, B. elata, y B. annua. © grupo II lo forman: B. quiriegoensis, B. eludens, B. aristidoides var. aristidoides, B. barbata var. barbata, B. eriopoda, B. radicosa, B. barbata var. rothrockii, B. williamsii, B. barbata var. sonorae, B. eriostachya, B. reflexa, B. breviseta, B. gracilis, y B. repens. Mientras que el grupo III esta integrado por B. hirsuta var. glandulosa, B. rigidiseta, B. chasei, B. alamosana, B. hirsuta var. hirsuta, B. scorpioides, B. chondosioides, B. curtipendula var. caespitosa, B. johnstonii, B. simplex, B. curtipendula var. curtipendula, B. triaena, y B. karwinskii. Como puede verse el dendrograma resultante, en el andlisis agrupa en general las mismas especies, con algunas diferencias irrelevantes. El dendrograma realizado en este trabajo muestra por un lado que todas las especies comparten caracteres comunes entre ellas como son, la forma de los cuerpos de silice, las ondulaciones en las paredes horizontales de las células largas con sinuosidades en forma de Q, los micropelos bicelulares, forma de los estomas, etc. Se encontraron diferentes patrones en el arreglo y ntimero de las hileras de células cortas-células largas en la zona costal: 1) zona costal del centro mds ancha que el resto, 2) tercera y cuarta zona costal de ambos extremos de la lamina mds ancha que las demas, 3) intercalandose zonas costales anchas y angostas, y 4) zonas costales homogéneas en toda la lamina. Rosales-C. & Herrera-A.: Bouteloua epidermal anatomy 63 CONCLUSIONES De acuerdo con Metcalfe (1960) los caracteres anat6micos de la lamina de la hoja son significativos para la separaci6n de especies, de géneros 0 de taxas mayores de plantas. Con los resultados obtenidos en el presente trabajo, se puede concluir por un lado que las especies se delimitan perfectamente con base en sus caracteres anat6micos de epidermis, ejemplo de ello es el caso de Bouteloua ramosa que no present6 diferencias significativas con B. breviseta, de la cual parece ser sindnimo. Por otra parte el andlisis fenético justifica la separacién de algunas especies en variedades ya que al observar los grupos del dendrograma estas variedades quedan alejadas entre si ya sea en el mismo grupo o en grupos diferentes. Por otra parte, existen discrepancias en la validéz del nivel genénco de Bouteloua, como una unidad, o si debe ser separado en dos géneros (Bouleloua Lag. y Chondrosum Desv.) como lo proponen algunos autores: Clayton & Renvoize (1986); Davidse, Sousa, & Chater (1994). Mientras que autores como: Gould (1979); McVaugh (1983); y Beetle, ef al. (1987), consideran subgéneros a Chondrosum y Bouteloua; incluyendo en el primer subgénero a las especies conocidas comunmente como Navajitas y en el segundo subgénero a todas las especies conocidas comuinmente como Banderillas. Con base en caracteres anat6micos de la epidermis, en el presente trabajo se llega a la conclusién de apoyar la delimitaci6n de un solo género, por los patrones comunes que presentan todas las especies. Sin embargo se debe esperar a conjuntar los resultados de los diferentes caracteres que se estudian del género Bouteloua, para \legar a una conclusi6n mejor fundamentada. En el dendrograma resultante del andlisis fenético de este estudio, las especies se agrupan de forma diferente a como se agrupan tomando en cuenta los caracteres morfoldgicos unicamente (Herrera & De la Cerda 1997), de donde se percibe que los grupos de caracteres morfoldgicos y anat6micos no se comportan por igual entre las especies, concluyendose que posiblemente la evoluci6n anat6mica de la epidermis, durante la especiacién del género Bouteloua, no siguid los mismos causes que la morfolégica. Puesto de otra manera, puede interpretarse como que los cambios anat6micos de la epidermis no fueron influenciados por los mismos factores ecofisiograficos que defienieron las diferentes especies y variedades. La homogeneidad en los caracteres anat6micos de la epidermis y los grupos de especies formados en el dendrograma muestra que éstos pudieron evolucionar por separado de los caracteres morfoldégicos, ya que los grupos no se forman solamente de especies conocidas comtinmente como banderilla 0 como navajitas de acuerdo a la forma de la inflorescencia, estos dos grupos se mezclan entre ellos debido a las caracteristicas anat6micas comunes. Como conclusién adicional es importante sefialar que los estudios de la anatomia de la epidermis son importantes a nivel genérico, ya que pueden ayudar a reconocer facilmente diferentes géneros de gramineas cuando no se cuente con estructuras morfol6gicas suficientes para la determinaci6n de tales grupos. 64 PHYTOLOGIA July 1996 volume 81(1):46-64 AGRADECIMIENTOS Se agradece el financiamiento al presente estudio del IPN, UAA, CONABIO- BO61 y CONACyT- 3098-N. Asi también se agradece a los curadores de los Herbarios ANSM, CIIDIR, ENCB, HUAA, IBUG, IEB, MEXU, SLPM, y UAG; por las facilidades recibidas en la consulta de los ejemplares. A Margarita de la Cerda, gracias por sus sugerencias y correcciones al presente manuscnito. LITERATURA CITADA Beetle, A.A., E. Manrique F., V. Jaramillo L., P. Guerrero S., A. Miranda S., Nufiez T., & A. Chimal H. 1987. Las Gramineas de México. Tomo II. Comisién Técnico Consultiva para la Determinacién de Coeficientes de Agostadero, Secretarfa de Agricultura y Recursos Hidraulicos, Ciudad México, México. Clayton, W.D. & S.A. Renvoize. 1986. Genera Graminum, Grasses of the World. 389 pp. Her Majesty's Stationery Office., London, Great Britain. Davidse, G., M. Sousa S., & O. Chater A. 1994. Flora Mesoamericana, Alismataceae a Cyperaceae. UNAM, Missouri Botanical Garden. Vol. 6, 543 pp. Ebinger, E.J. & L.J. Carlen. 1975. Culm morphology and Grass Systematics. Eastern IIlinois University, vol. 68(2):87-101. Ellis, R.P. 1979. A procedure for standardizing comparative leaf anatomy in the Poaceae. II. The epidermis as seen in surface view. Bothalia 12(4):641-671. Gould, F.W. 1979. The Genus Bouteloua (Poaceae). Ann. Missoun Bot. Gard. 66(3):348-416. Griffiths, D. 1912. The Grama Grasses: Bouteloua and related Genera. Contr. of the U.S. Natural Herbanum. 14(3):343-428. Herrera-A., Y. & M. De la Cerda-L. 1997. Morphometrics of the mexican Bouteloua. Phytologia. In press. Kovach, W.L. 1987. A Multivariate Statistical Package. Ver. 1.31., Department of Biology. Indiana University, Bloomington, Indiana. McVaugh, R. 1983. Flora Novo-Galiciana, Vol. 14, Gramineae. University of Michigan Press, Ann Arbor, Michigan. Metcalfe, C.R. 1960. Anatomy of the Monocotyledons. |. Gramineae. 731 pp. Clarendon Press, Oxford, Great Bnitain. Phytologia (July 1996) 81(1):65. INDEX TO AUTHORS, VOLUME 80 Arreguin-Sanchez, M.L. 8, 329 Robinson, H. 350 Athar, M. 385 Singhurst, J.R. 30, 280 Averett, J.F. 273 Siqueiros Delgado, M.E. 108 Babutina, K.A. 48 Turner, B.L. 92, 95, 100, 115, 118, Basinger, M.A. 352 128, 133; 145,253; 257; 276, Borowski, M. 30 285, 291, 368 Carter, R. 288 Vicente O., J.A. 40 Chantaranothai, P. 140 Volz, P.A. 48 D’Arcy, W.G. 273 Warnock, M.J. 148 Esparza Sandoval, S. 73 Wipff, JK. 35, 288, 343, 348 Galan de Mera, A. 40 Wivagg, D.E. 23 Gooch, J.R. 280 Worthington, R.D. 121 Herrera Arrieta, Y. 73 Hinton, G.S. 58, 62 Holmes, W.C. 23, 30, 280 Jones, S.D. 288 Little, R.J. 295 MacRoberts, B.R. 1 MacRoberts, M.H. 1 Morgan, T.L. 280 Palacios-Chavez, R. 8, 329 Patterson, T.F. 104 Pryor, D.R. 296 Quiroz-Garcia, D.L. 8, 329 Reed, C.F. 284 Riefner, Jr., R.E. 296 Robertson, P. 352 65 Phytologia (July 1996) 81(1):66. INDEX TO REVIEWERS, VOLUME 80 The editor expresses his most sincere appreciation to the following individuals. These are persons who have reviewed papers that were submitted for publication in volume 80 of Phytologia. Without the willingness and diligence of these reviewers, the task of the editor would be much more difficult, and the quality of the papers published would be lessened. To each of you, I offer my most sincere thanks. Michael J. Warmock, Editor Allen, C.M. Mayfield, M. Bartholomew, B. McDaniel, S. Bowler, P. Nesom, G.L. Boyd, S. Newman, M. Bretz, W. Phillipe, L.R. Buchfinck, E.A. Pruski, J. de la Cerda, M.C.M. Robinson, H. Delevoryas, T. Rodriguez J., C. Evans, R.E. St. John, T. Fernandez N., R. Shlemon, R. Fox, III, W.E. Simpson, D. Fryxell, P.A. Siqueiros, M.E. Hatch, S.L. Thomas, R.D. Herrera Arnieta, Y. Todzia, C. Hunter, R.L. Tovar S., O. Jones, C. Tumer, BL: Jones, G.D. Turner, G. Jones, S.D. Ulaszek, E. Kuhn, G. Wendt, T. Lellinger, D.B. Williams, J. MacRoberts, D.T. Wright, D. Mateus P., C.L. Wurdack, J.J. Phytologia (July 1996) 81(1):67 Issue 80(1) 80(2) 80(3) 80(4) 80(5) 80(6) PUBLICATION DATES, VOLUME 80 Cover date January 1996 February 1996 March 1996 Apnil 1996 May 1996 June 1996 67 Publication date 27 September 1996 31 October 1996 10 December 1996 23 December 1996 4 February 1997 21 February 1997 NOW AVAILABLE, PHYTOLOGIA MEMOIRS, Volume 11 THE COMPS OF MEXICO, Volume 1 EUPATORIEAE B.L. TURNER 1997, 8 1/4 x 10 3/4 in., 272 pp., illus., $44.95, ISBN 0-9642085-2-0. The second of an anticipated ten volume series on the Asteraceae of México, this volume includes the tbe Eupatonieae. Other volumes will include the remainder of the 2700+ species of Asteraceae known from México. The Mexican representatives of the family are particularly rich in the tribes Helenieae, Heliantheae, and Eupatorieae. 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