_PHYTOLOGIA is ; . a An international journal to expedite plant systematic, wii eel ac : ie Rg gets and ecological pecan Vol. 80 Pe Apa mye 2 No. 4 = cS ee eke oles _ TURNER, B. sg New. species and combinations in’ Pseudogynoxys i (Senecioneae)... Phriday Aad ohiae's Thu Se LeimTv RUT geod Coie Aiea NE nee ees 253 oe ‘TURNER, B.L., Revision of Dieamanehouein (Asteraceae). o2o.0. 0. ee, 257 a _D'ARCY, W.G. & J.B. “AVERETT, Recognition of tribes Capsiceae and fee ‘Physaleae, subfamily Solanoideae, Solanaceae... 273 Bi a TURNER, B.L., A new Species . of Roldana (Senecioneae) from Oaxaca, NEE SG = RRR aE PeOn. Aa ee US eR Me ag et aes Se Se a le 276 Ee HOLMES, W.C., PL: MORGAN. IR GOOCH, & IR. SINGHURST. ak Comments on the distribution of Botrychium lunarioides (Ophioglossaceae) LOBES RIES TRIE ANS Cag acl herd A eg ape CDE Niger pe eee 280 a REED, boy, ake New combinations for the flora of the central eastern United as SRC 0 2 APIS SaE ius EH ce oO Lag eo Ae CGS RE 284 - es om _ TURNER, B. Toe * new. species of Golan (Ribiadee) from northeastern «PSNI SS 1s Fore alt ME aaa RE ic eS Se ag a a . £285 oe fs JONES, Ss. D., J. ia “WIP, & R. CARTER, Nomenclahiral combinations in “LIBRARY. SAN uf 1997 ‘NEW YORK _ _ BOTANICAL GARDEN ore ~ Published iy Michael J. Watsbek Bik ad ed ae 185 5 Westie Drive Huntsville, Texas 77340 USA. ae TecaGhh. is. elmeiech on acid free Saini ne * PHYTOLOGIA An international jourial to expedite plant systematic, phytogeographical and ecological py tien NORD ae April 1906 2 No. 4 CONTENTS . TURNER, B. gr - New. SPORES and combinations in Pseyudogynoxys. : (Senecioneae) Sena e Nene teete | ee CaS lei Se gUR Ty ed Gaahed Se Gan Tee oxen e Bad tisaas 253 .» TURNER, B.L., Revision of Desmanthodium (Asteraceae). 222.20... 257 et D’ARCY, W.G. & LE. AVERETT, Recognition of tribes Capsiceae and | Physaleae, subfamily Solanoideae, Solanaceae. ........00. 050. 273 Be TURNER, B.L., A new species of Roldana (Senecioneae) from Oaxaca, 5; t AMS 2 pte Ric 2 spss ean Loren RE NG Fe one eh S Le ie AV re ep ae eG eae ae 276 x HOLMES, W.C., T.L: MORGAN, J-R.. GOOCH, & LR. SINGHURST, _.» Comments on the distribution of Botrychium lunarioides' (Ophioglossaceae) A Ee BS pte tate eae cae eeepc IAN Ger ti me Mears eS Nee ort PE AL SH 280 REED, GOP. : ‘New combination for the flora of the central eastern’ United Dip eG pen aig SOM aU ONS Wi lanstieg nots CFE ge URE ela watt pee aun ce mtg ese 284 oe TURNER, Biles oe new. species of Calan (Rubiaceae): from northeastern Ag WARDMAN He eet eos hide Peete ari eed oe RETO Fi aD Te 285 rie JONES. S.D., J.K. WIPER, & R. CARTER, Nomenclatural combinations in Cyperus (Cyperaceae). Bis ae Nain Gor EOE oe acts pe bPOR Recados eae 288 _ TURNER, B. L., A new vancly: of ‘Coreopsis tnutica (Asteraceae) from western ee iadene Ge a eu ee ree Ve aes 291 « LITTLE, RJ; SRoniencatersl correction in Viola (Violaceae). ..... Satie ban 295 RIEFNER, Ir., R.E. & D.R: PRYOR, New locations and interpretation of Ae vernal pools in southern California. BS Seo Ne a Le RS a a 296 aM -Phytologia Memoirs available etree chan Aas 3 SPST mer Sey aae orn overt ey 328 eh Bact oF / 1 S BOTANICAL GARDEN — “Published ii Michael ce Waenack ” 185 5 Westridge Drive Huntsville, Texas 77340 U.S-A. EOC: is- een on acid free paper. anes: re a ; die taicit d ’ it + Se Fes Phytologia (April 1996) 80(4):253-256. NEW SPECIES AND COMBINATIONS IN PSEUDOGYNOXYS (SENECIONEAE) B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A new species of Pseudogynoxys s.l., P. alajuelana B.L. Turner, is described from Costa Rica. It belongs to a group of species that some workers would include in or near the recently described genera, Garcibarrigoa Cuatr. and Talamancalia H. Rob. & Cuatr., elements of which I position in an expanded Pseudogynoxys. To this end the following new combinations are proposed: Pseudogynoxys westonii (H. Rob. & Cuatr.) B.L. Turner, comb. nov., and P. durandii (Klatt) B.L. Turner, comb. nov. So construed, Pseudogynoxys is conceived as a mesophytic genus of clambering shrubs and/or perennial herbs having mostly orange ray florets and variable style branches, the appendages of the latter varying from apically rounded and penicillate basally, to long-attenuate and sparsely penicillate along their margins. KEY WORDS: Asteraceae, Senecioneae, Pseudogynoxys, Garcibarrigoa, Talamancalia, Costa Rica, systematics Pseudogynoxys was first proposed by Greenman (1902) as a subgenus of Senecio. Cabrera (1950) elevated this to generic rank, the group largely recognized by its clambering or scandent habit, orange rays, and especially by its style branches which were thought to be rather uniformly bestowed with triangular stylar appendages fringed at the base with a crown of hairs. Since Cabrera’s treatment, several newly described genera (Garcibarrigoa Cuatr. and Talamancalia H. Rob. & Cuatr.) have been proposed that clearly relate to Pseudogynoxys, differing from the latter primarily in habit (non-clambering perennial herbs) and stylar appendages. In my view (Tumer 1991), Garcibarrigoa and Talamancalia are readily positioned in an expanded concept of Pseudogynoxys, unless one prescribes to a microgeneric concept for the numerous tropical American elements of Senecio as proposed by Nordenstam & Pruski (1995), Robinson & Cuatrecasas (1994) and perhaps others (Barkley, pers. comm.). Much of the reasoning behind the generic splintering of the pseudogynoxoid alliance referred to above has to do with emphasis upon microcharacters, most notably 253 254 PHY TOLOGIA Apnil 1996 volume 80(4):253-256 stylar branches and their appendages. Thus Pruski (1996), in his defense of the retention of Garcibarrigoa, notes that species of the latter “have neither the scandent habit nor the sterile triangular to acuminate style branch appendages tipped by a fringe or tuft of papillae, as typical of Pseudogynoxys.” While his habital observations are correct, my survey of style branches in the pseudogynoxoid alliance show a wide range of variation in this character, even within Pseudogynoxys (s.s.). For example, elongate stylar appendages with varying degrees of basal hairs, to those with merely rounded apices which are minutely papillate at base occur in Pseudogynoxys (s.s.). We (Mendenhall & Turner, in prep.) are currently undertaking a scanning electron microscopical study of the stylar appendages of the pseudogynoxoid complex and preliminary results suggest that stylar characters in this group are much more plastic than heretofore supposed, and that treatment of these taxa within an enlarged genus Pseudogynoxys will prove defensible on phyletic grounds. PSEUDOGYNOXYS ALAJUELANA B.L. Turner, spec. nov. TYPE: COSTA RICA. Alajuela: P. Nac. Rinc6én de la Vieja. Quebrada Leiva. Colonia Blanca (10° 47’ 20” N, 85° 15’ 20” W), 600 m, 4 Apr 1991, Gerardo Rivera 1235 (HOLOTYPE: MO!). Similis P. boquetensis (Standl.) B.L. Turmer sed habens capitula sine floribus radiantibus, appendices styli lineares-lanceolatas, et lobos corollarum faucibus ca. 1/3 plo longiores. Perennial herbs 35-40 cm high. Stems densely hirsute with crinkly, purplish- septate, hairs. Leaves simple, gradually reduced upwards, those at mid-stem 8-10 cm long, 3-4 cm wide; petioles 2.0-2.5 cm long, auriculate at the base, subclasping but not connate; blades elliptic, pinnately veined, densely pubescent above and below, the margins irregularly serrate. Heads 2-4, arranged in loose corymbs, the ultimate peduncles mostly 3-10 cm long. Calyculus of 10-14 loosely arranged linear bracts 3-5 mm long. Involucre ca. 1 cm long, composed of ca. 11 lanceolate bracts. Ray florets absent. Disk florets 80-100 (est.); corollas tubular, glabrous, reportedly yellow, ca. 12 mm long, the throat poorly defined, 3-4 mm long, the lobes 5 ca. mm long, narrowly deltoid. Anthers ca. 2 mm long, basally obtuse or rounded, the apical appendages ca. 0.5 mm long. Style branches ca. 4 mm long, the appendages linear- lanceolate, hispidulous. Achenes (immature) 3-4 mm long, columnar, 8-10 costate, minutely appressed-pubescent; pappus of numerous delicate, white, readily deciduous bristles 10-12 mm long. Pseudogynoxys alajuelana is an enigmatic species, possessing corolla and stylar characters which are seemingly attenuated forms of typical elements of the genus, but having the habit of those species that Robinson & Cuatrecasas (1994) recognize as the segregate genus Jalamancalia. The corolla lobes of P. alajuelana are narrowly trianguloid and ca. 1/3 as long as the throat; the stylar branches have appendages that are linear-lanceolate and lack a well-defined basal tuft of hairs. Among most species of Pseudogynoxys, corolla lobes are narrowly triangular and usually 1/2 or more as long as the tube; the stylar appendages are usually narrowly deltoid to narrowly lanceolate, with often well-defined basal hairs. Nevertheless there is much variation in these two characters among the species | have examined, especially in the stylar appendages (as noted above) which may vary from merely trianguloid to lanceolate Tumer: New Pseudogynoxys 255 with varying degrees of pubescence, occasionally within a single head. Regardless, I can not believe that the short corolla lobes and linear-lanceolate, weakly penicillate stylar branches of P. alajuelana are anything but accentuations of trends already present in Pseudogynoxys s.s.. The habit of P. alajuelana, a perennial non- clambering herb ca. 40 cm high, to judge from the single specimen examined, is that found in the recently proposed Talamancalia (Robinson & Cuatrecasas 1994). Additionally, Pseudogynoxys alajuelana has leaves similar to Talamancalia boquetensis (Standl.) H. Rob. [= Pseudogynoxys boquetensis (Standl.) B.L. Turner], possessing subclasping basal appendages, (as do some species of Pseudogynoxys, as noted by Robinson & Cuatrecasas 1994). Various authors might possibly consider the present novelty to be yet another closely related monotypic genus, if emphasis is placed upon the stylar appendages and corolla lobes possessed by Pseudogynoxys alajuelana, for Robinson & Cuatrecasas (1995) note that The five genera, Garcibarrigoa, Jacmaia, Jessea, Pseudogynoxys, and Talamancalia, differ from almost all species of Senecio sensu stricto by having long and narrow lobes on the corolla. The lobes in Jacmaia and all of Jessea except the type species are about as long as the throat. The limbs and the lobes of the corolla of Garcibarrigoa are comparatively short with the lobes narrowly tnangular rather than narrowly oblong. My own view is that the habit and short corolla lobes of Pseudogynoxys alajuelana Vitiate recognition of a monotypic Garcibarrigoa. Indeed, I believe that phyletic studies emphasizing similarities (instead of differences) among the several taxa mentioned in the above will show that Pseudogynoxys, Garcibarrigoa, and Talamancalia are more closely related among themselves than they are to Jacmaia and Jessea. DNA studies are sorely needed to help ferret out relationships among these various generic segregates. In line with the above taxonomic views | propose the following new combinations: PSEUDOGYNOXYS WESTONII (H. Rob. & Cuatr.) B.L. Turner, comb. nov. BASIONYM: Talamancalia westonii H. Rob. & Cuatr., Novon 4:52. 1994. PSEUDOGYNOXYS DURANDII (Kliatt) B.L. Turner, comb. nov. BASIONYM: Senecio durandii Klatt, Bull. Soc. Bot. Belg. 31:211. 1892. This Costa Rican endemic is a suffruticose herb with pendant branches having orange ray and disk florets and emits “an unpleasant odor when crushed” (Almeda 5791 [TEX]; described as that of “culantro” by Grayum 3746 [TEX]). Standley (1938) notes the species to be “a most distinct one, altogether unlike any other with which I am familiar.” He also adds that the plant appears to be rare occurring “on rocks at the edge of streams in deep, dark forests, sometimes in the spray of waterfalls.” This is also borne out by label data on the ten or more sheets known to me (LL, MO, TEX). 256 PHY TOLOGGIA Apmil 1996 volume 80(4):253-256 Of the species to be included within my treatment of Pseudogynoxys for Costa Rica (in prep.) this is the most distinctive, but it appears to me to fall within the pseudogynoxoid alliance as conceived here. ACKNOWLEDGMENTS I am grateful to Gayle Tumer for the Latin diagnosis, and to Ted Delevoryas for reviewing the manuscript. Maria Thompson provided the illustration. LITERATURE CITED Cabrera, A. 1950. Notes on the Brazilian Senecioneae. Brittonia 7:53-74. . Greenman, J.M. 1902. Monographie der nord- und centralamerikanischen Arten der Gattung Senecio. 32:1-33. Nordenstam, B. & J.F. Pruski. 1995. Additions to Dorobaea and Talamancalia (Compositae: Senecioneae). Compositae Newsletter 27:3 1-42. Pruski, J.F. 1996. Pseudogynoxys lobata (Compositae: Senecioneae), a new species from Bolivia and Brazil. Syst. Bot. 21:1-4. Robinson, H. & J. Cuatrecasas. 1977. Notes on the genus and species limits of Pseudogynoxys (Greenm.) Cabrera (Senecioneae, Asteraceae). Phytologia 36: 177-192. Robinson, H. & J. Cuatrecasas. 1994. Jessea and Talamancalia, two new genera of the Senecioneae (Asteraceae) from Costa Rica and Panama. Novon 4:48-52. Standley, P.C. 1938. Flora of Costa Rica. Publ. Field Mus. Nat. Hist. Vol. 18. 1616 pp. Turner, B.L. 1991. Transfer of two species of Senecio to Pseudogynoxys (Asteraceae - Senecioneae). Phytologia 71:205-207. Phytologia (Apnil 1996) 80(4):257-272. REVISION OF DESMANTHODIUM (ASTERACEAE) B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT The genus Desmanthodium (Asteraceae, Heliantheae) is treated as having seven species, six of these indigenous to México and Central America, and one to South America. The recently described D. congestum Armiagada & Stuessy, is believed to be the same as D. tomentosum. A key to the species, selected illustrations, and complete synonymy are provided, along with dot maps showing their distribution. KEY WORDS: Asteraceae, Heliantheae, Desmanthodium, México, systematics The present study has been occasioned by my treatment of the tribe Heliantheae for the Asteraceae of Mexico (cf, Turner 1996). As conceived here, Desmanthodium is a small mostly montane genus of seven species, six of these native to México. The species are very closely related, and if partitioned into infrageneric components they would seem to fall into two groups: a strictly herbaceous element, D. ovatum Benth., with the remainder forming a subclade of robust suffruticose herbs, subshrubs or shrubs to 4 m high. Desmanthodium was first proposed by Bentham in 1876 with the description of two Mexican species, D. perfoliatum Benth. and D. ovatum, the former subsequently selected as the generitype. These were positioned by Bentham (Genera Plantarum, 1876) in the tnbe Heliantheae, subtribe Milleninae, between the genera Riencourtia and Clibadium. Thereafter, among the species recognized here, Hemsley added D. guatemalense Hemsl. in 1881; Greenman proposed D. fruticosum Greenm. in 1903; T.S. Brandegee described D. tomentosum T.S. Brandegee in 1914; Blake in 1924 conjured up D. blepharodon S.F. Blake, and Turner concocted D. hintoniorum B.L. Turner in 1996. A few other names have been proposed, but these have been treated as synonyms. There has been no revision of the genus prior to the present study, although Robinson (1981) provided a succinct account of its subtribal position, as noted under comments presented below (cf, Generic Relationships). cot 258 PHY TOLOGIA Apnil 1996 volume 80(4):257-272 CHROMOSOMES As indicated in the list of chromosome counts tabulated below, all counts were determined from meiotic material. Fay (1974) was the first worker to report a chromosome number for Desmanthodium, this being a count of n=18 pairs for D. fruticosum. Subsequent workers have confirmed this number for the genus, except for Ralston, et al. (1989) who report a count of n=17 pairs for D. fruticosum. The latter count should be confirmed since it does not agree with previous reports and was not documented by photographs or camera lucida drawings. In fact, none of the counts in Desmanthodium has been documented by photographs or illustrations. CHROMOSOME COUNTS FOR DESMANTHODIUM D. fruticosum 2n=36 Fay (1974) Oaxaca: Cronquist 10855 D. fruticosum 2n=36 Keil & Stuessy (1977) Mexico: Stuessy 3129 D. fruticosum 2n=36 Keil, et al. (1988) Guerrero: Keil 15356 D. fruticosum 2n=34 Ralston, et al. (1989) Guerrero: Turner 15876 D. perfoliatum 2n=36 Sundberg, ef al. (1986) | Oaxaca: Turner 80A (reported as D. caudatum) GENERIC RELATIONSHIPS Robinson (1981) recognized Desmanthodium, along with Stachycephalum, as the only two genera in his subtribe Desmanthodiinae, numbered 6 from among 32 subtribes in his breakdown of the tnbe Heliantheae. Desmanthodium differs markedly from Stachycephalum in having its ray florets completely enclosed in a sac or vesicle, in addition to yet other characters discussed by Robinson. He reckoned the Desmanthodiinae to be closely related to his subtribe number 7, the Clibadiinae, which contained three genera (Clibadium, Lantanopsis, and Riencourtia). Nearly all workers would agree that the two subtnbes recognized by Robinson are highly specialized. Indeed, Robinson (1981, p. 40) conjectures that the so called vesicle which houses the ray florets in Desmanthodium might actually be a loosened part of the ovary wall, as opposed to an enveloping involucral bract or phyllary; so interpreted this would be a unique feature in the Asteraceae. Whether or not Stachycephalum is properly positioned with Desmanthodium in the subtribe Desmanthodiinae is moot; it might with equal morphological justification be included with or near Clibadium or Lantanopsis in the subtribe Clibadiinae. The probable base chromosome number of Desmanthodium (x=18) differs from that of Clibadium (x=16, cf. Stuessy & Arriagada 1993) and Riencourtia (x=ca. 16); unfortunately chromosome counts for Stachycephalum are unreported. Turner: _ Revision of Desmanthodium 259 TAXONOMY DESMANTHODIUM Benth. Suffruticose stiffly erect perennial herbs to 1 m high, or erect to sprawling shrubs or subshrubs to 4m high. Leaves simple, opposite throughout, subpinnately nervate with usually 3 prominent nerves from above the base. Heads ill-defined, arranged in congested bracteate glomerules, the glomerules in tum disposed in very open cymose panicles, or relatively congested in flat-topped cymes. Involucral bracts 1-3, separate to the base, not forming a clearly defined involucre. Receptacle plane, epaleate, except for an outer series of pales which completely enwrap the subtended ray florets. Ray florets 1-3, pistillate, fertile, enclosed in sac-like bracts; corollas ca. 1 mm long, the ligule absent or nearly so. Disk florets 5-10 per head, perfect but sterile, the ovaries elongating with age; corollas small, white, S-lobed; tubes about as long as or somewhat longer than the funnelform or campanulate throats. Acheres lenticular, black, minutely striate, completely enclosed in a persistent papery sac. Base chromosome number, x=18. Type species, Desmanthodium perfoliatum Benth. KEY TO SPECIES 1. Suffruticose herbs with slender simple stems 0.4-1.0 m high.......... 1. D. ovatum 1. Shrubs, or arching to sprawling thick-stemmed shrublets 1-4 m Pi see (2) 2. Leaves sessile, those at midstem clearly sen is bene D. perfoliatum eeeeeorves petiolate, never perfoliata 0.05.68 eh seer pe ag are tenes (3) 3. Stems and branches of the capitulescence glabrous; heads on thick stout peduncles, scarcely exceeding the foliage; Oaxaca, México Semen DAINMANAR) 2:5. ci n.cccns, (uel Rs Uses eee eros 3. D. hintoniorum 3. Stems and branches of the capitulescence clearly pubescent, either in lines or SaORA ERENCE. 5, ci) nied “elaade ls HOE SiN AIR: wince lect Meee Pave ee ee eee (4) 4. Uppermost stems and branches of the capitulescence tomentose throughout, the vestiture mostly 0.5-0.7 mm high; southwesternmost Chiapas, México and closely adjacent Guatemala: cai... cviicen Ren. We 5. D. tomentosum 4. Uppermost stems and branches of the capitulescence pubescent in lines, the vestiture mostly 0.2-0.4 mm high; widespread. ....................ccceeeeeeeeees (5) 5. Leaves on primary stems mostly thin, the blades 5-10(-15) cm long, decidedly ovate, widest near base or well below the middle (rarely not); Pacific montane slopes of western México from Durango to Oaxaca................. 4. D. fruticosum 5. Leaves on primary stems mostly thick (subsucculent), the blades (10-)15-20 cm long, elliptical, mostly widest at or near the middle; Guatemala, Honduras, Hl penseet ont Southokmericacincs.. 2 wok. A. BRR, ee (6) 6. Leaves on primary stems with blades gradually tapering, ciliate at the base, the petioles. 3-15. mm.long;, Venezuela.n...:..808.8n IR: 7. D. blepharodon 260 PHYTOLOGIA Apnil 1996 volume 80(4):257-272 6. Leaves on primary stems with blades not tapering upon the petioles, glabrous at the base, the petioles 1-5 mm long; Guatemala, Honduras and El Salvador. ....................065 6. D. guatemalense 1. DESMANTHODIUM OVATUM Benth., Hook. Icon. Pl., t. 1116, 1872. Fig. 1. TYPE: MEXICO. Oaxaca: “woods of the province of Oaxaca”, 7000-8000 ft, Nov-Apr 1840, Galeotti 2081 (LECTOTYPE [selected here]: K; Photoisolectotype: MICH!). Two collections were cited in the protologue, that selected here as lectotype and Andrieux 319 from the Sierra San Felipe, Oaxaca. Desmanthodium lanceolatum Greenm., Proc. Amer. Acad. Arts 34:576. 1899. TYPE: MEXICO. Morelos: mountains above Cuernavaca, 2100 m, 9 Aug 1898, C.G. Pringle 6940 (HOLOTYPE: MO; Isotype: UC’). Perennial glabrous herbs with simple, mostly unbranched stems 20-100 cm high; roots thickened, fasciculate or arising from short stout rhizomes (cf., Iltis 1289 [TEX]). Stems sparsely puberulent to glabrate, 2-4 mm thick below. Leaves well- spaced along the stem, mostly shorter than the internodes, those at midstem mostly 5- 15 cm long, 3-6 cm wide; petioles 5-35 mm long; blades ovate to subdeltoid, gradually or abruptly tapering upon the petioles, glabrous or nearly so. Heads arranged in relatively few, long-pedunculate aggregations. Outer and interior bracts of the aggregations ovate, subsucculent, their apices white or whitish. Heads 8-24 in any one aggregation, the whole superficially resembling a head (syncephalum), these borne upon peduncles 1-8 cm long. Heads with 1-3 pistillate florets and 12-16 sterile disk florets. Achenes flattened tangentially, ca. 3 mm long, 1.5 mm wide, encased in sac-like sparsely pubescent involucral bracts. DISTRIBUTION (Figure 5) AND ECOLOGY: México State, Morelos, Puebla and Oaxaca, mostly oak woodlands, 2000-2600 m, Jul-Aug. REPRESENTATIVE SPECIMENS: MEXICO. México State: Rzedowski 30918 (LL,MEXU). Puebla: Tenorio 7492f (TEX). Oaxaca: Panero 3617 (TEX); Pringle 4694 (LL,MEXU); Soule 2422 (MEXU,TEX). This species is well represented in herbaria and, because of its herbaceous habit, relatively easily recognized. Most collectors note the plants to be between 20-100 cm high; however, label data on Hinton 2758 (LL) records the plant to be 1.5 m high, but this is not evident from the sheet concerned. I am unable to distinguish Desmanthodium lanceolatum from D. ovatum. The former name has been applied to plants having leaf forms with lanceolate to narrowly ovate blades which taper upon the petioles. Both leaf forms may occur in the same general region and intergrades between such forms occur. Exceptionally petiolate blades of D. ovatum also occur (Figure 1), but most collections lie somewhere pooh iin these extremes, the blades at midstem usually gradually tapering upon the petioles. One might consider erecting a monotypic section to house Desmanthodium ovatum, for it is sufficiently distinct from the shrubby elements of Desmanthodium to perhaps warrant such recognition. Its habit, relatively few aggregations to a capitulescence and generally more numerous staminate florets are diagnostic. Turner: Revision of Desmanthodium Figure 1. Desmanthodium ovatum (Panero 3617). 261 262 PHYTOLOGIA Apni 1996 4 f » Fe Wy) a? ¢, , Ahi x Jaf y ~ptee y we: A) AC Otay Figure 2. Desmanthodium perfoliatum (t. 1116, 1887). volume 80(4):257-272 Turner: - Revision of Desmanthodium 263 Figure 3. Desmanthodium hintoniorum (holotype). 264 PHYTOLOGIA Apni 1996 volume 80(4):257-272 Figure 4. Distribution of Desmanthodium fruticosum (open circles); D. guatemalense (open triangles); D. hintoniorum (closed triangle); D. tomentosum (closed circles). 265 Revision of Desmanthodium Tumer. e sly ‘ ‘ ‘ 4 ay VENEZUELA s 4 ,, ‘ Mba! ®@eeen marina ; Inset, D. blepharodon (South Figure 5. Distribution of Desmanthodium ovatum America). 266 PHYTOLOGIA Apni 1996 volume 80(4):257-272 2. DESMANTHODIUM PERFOLIATUM Benth., Hook. Icon. Pl. 12:15. t. 1116. 1887. TYPE: MEXICO. Oaxaca: w/o specific locality, 4500 ft, 1835-1840, H.G. Galeotti 2050 (HOLOTYPE: K). Flaveria perfoliata Kliatt, Leopoldina 23:146. 1887. TYPE: MEXICO. Oaxaca(7): “Cumbre de Estepe”, 1841-43, F.M. Liebmann 482 (LECTOTYPE [selected here]: GH!). Rydberg (1915) was the first to call attention to the synonymy listed here. Desmanthodium caudatum S.F. Blake, J. Wash. Acad. Sci. 28: 488. 1938. TYPE: MEXICO. Chiapas: Escuintla, Finca Juarez, 12 Aug 1937, E. Matuda 1750 (HOLOTYPE: US!; Isotype: MEXU; Fragment holotype: LL!). Shrublets or shrubs 1-4 m high. Much resembling Desmanthodium guatemalense but the leaves relatively thin and clearly perfoliate, the blades united below, not at all petiolate, and heads more numerous; chromosome number, 2n = 36. DISTRIBUTION (Figure 6) AND ECOLOGY: Guerrero, Oaxaca and Chiapas in pine-oak forests, 900-1700 m; Jul-Nov. REPRESENTATIVE SPECIMENS: MEXICO. Guerrero: 15.8 mi by road from Chilpancingo west towards Omiltemi, pine-oak forest, 2000 m, rare among limestone boulders, 27-28 Jul 1968, Anderson 4935 (MICH). Oaxaca: Sierra San Felipe, 7500 ft, 13 Aug 1894, Pringle 4799 (MICH); 3.6 mi SW of Suchixtepec along road to Puerto Angel, 22 Aug 1980, Turner 80A-11 (TEX). Chiapas: Mpio. Bochil, 4 mi NE Bochil, 4500 ft, 21 Aug 1965, Breedlove 12073 (LL,MICH); ca. 16 mi W of San Cristébal de las Casas, 26 Aug 1976, Hartman 4179 (TEX); Mpio. Amatenango del Valle, 5900 ft, 5S Sep 1966, Ton 1104 (LL). Like Desmanthodium fruticosum, D. perfoliatum is a highly variable, relatively common species, especially in Chiapas. Only a single collection is known from Guerrero (cited above) where it was reported to be “Rare, seen once.” The plant concerned has relatively narrow thin leaves (on secondary shoots), the blades markedly narrowed below with weakly developed auriculate-perfoliate bases. Specimens identified as D. caudatum are robust with very large, relatively thin leaves, but similar leaf forms occur across the range of the species (e.g., in Oaxaca, Turner 80A-11, cited above), and there seems little justification for its recognition. 3. DESMANTHODIUM HINTONIORUM B.L. Turner, Phytologia 79:317. 1996. Figure 3. TYPE: MEXICO. Oaxaca: Mpio. Miahuatlan, La Sirena, 2525 m, 23 Oct 1995, Hinton, et al. 26409 (HOLOTYPE: TEX!). Shrub to 1.5 m high, the stems clearly woody and glabrous throughout. Leaves mostly 10-12 cm long, 3.0-3.5 cm wide; petioles 2-4 mm long; blades narrowly elliptical, pinnately nervate, gradually tapering upon the petioles, the margins with minute well-spaced, denticulate teeth, but seemingly entire upon superficial inspection. Heads much-congested and terminal on stout peduncles 0.5-2.0 cm long, the syncephalous structure ca. 1.5 cm high and 2-3 cm across. Bracts ovate, glabrous, subcoriaceous, 8-10 mm long, 5-6 mm wide, not forming a well-defined involucral- bound head. Receptacle plane, glabrous. Pistillate florets 2, fertile; ligule absent, the tube ca. 1.5 mm long; achenes ellipsoid, glabrous, completely enclosed in fused, elliptical (in outline) bracts, the latter 6-7 mm long, ca. 2.5 mm wide, glabrous Turner: . Revision of Desmanthodium 267 throughout. Disk florets ca. 8, sterile, the style branches fused, forming a conical brush ca. 2 mm long; corollas white, glabrous, 5-lobed, the lobes ca. 1.4 mm long with ill-defined veins, these scarcely marginal, if at all; base of style surrounded by a well defined nectary ca. 0.75 mm high. Achenes of disk florets elongating at anthesis up to several times their bud-size so as to resemble stout stalks 5-10 mm long. DISTRIBUTION (Figure 4) AND ECOLOGY: known only from type material. As noted in the onginal description, this taxon is closely related to the more southern, Desmanthodium guatemalense; it differs in having narrower, more elliptical, nearly entire leaves, and being glabrous throughout, including all floral parts. 4. DESMANTHODIUM FRUTICOSUM Greenm., Proc. Amer. Acad. Arts 40:37. 1904. TYPE: MEXICO. Jalisco: Zapotlén, 9 Oct 1903, E.W.D. Holway 5137 (HOLOTYPE: GH!). Sprawling or arching shrubs or subshrubs 0.5-4.0 m high. Young stems pubescent in lines with crinkly brownish or tannish hairs, the vestiture mostly 0.3 mm high or less. Midstem leaves (of primary shoots) mostly 8-17 cm long, 3-7 cm wide; petioles 3-10 mm long; blades ovate to ovate-elliptic, sparsely appressed-pubescent beneath along the major veins, the margins variously dentate. Heads aggregated, the aggregations arranged in terminal rounded corymbose panicles 15-30 cm high and about as wide. Involucres subtended by 1-3 leathery, whitish, broadly ovate bracts, at anthesis mostly 4-6 mm long, 2-4 mm wide. Ray florets 1, pistillate fertile. Disk florets 4-8, the corollas white, 2-4 mm long, sparsely pubescent to nearly glabrous; lobes ca. 0.8 mm long. Chromosome number, 2n =36. DISTRIBUTION (Figure 4) AND ECOLOGY: Western México from Durango to Oaxaca, occurring in pine-oak woodlands, 1800-2400 m; Aug-Oct. REPRESENTATIVE SPECIMENS: MEXICO. Durango: “near summit on Durango Road,” 10 Oct 1955, Templeton 7643 (MICH). Nayarit: ca. 10 road mi E of Jacocotlan, on road to Tepic, 4 Oct 1952, McVaugh 13360 (MICH). Jalisco: ca. 10 mi SSE of Autlan, 29 Sep 1960, McVaugh 19548 (LL,MICH). Colima: 22 km NNW of Colima, Rancho El Jabali, 26 Aug 1988, Sanders 8366 (TEX). Michoacdn: Mpio. Coalcoman, 15.1 mi SW of Coalcomdan, 12 Sep 1985, Luckow 2915 (TEX). México: 12.5 mi SW of Temascaltepec, 12 Oct 1966, Anderson 3945 (MICH). Guerrero: 62 road mi N of Acapulco, 20 Oct 1962, Cronquist 9706 (MICH,NY,TEX). Oaxaca: ca. 10 mi N of Putla, 30 Oct 1970, Cronquist 10855 (MICH,NY). This species is represented in herbaria by numerous collections and is nicely illustrated by McVaugh (1984). The foliage is quite variable, leaves on secondary shoots being smaller and narrower than those on primary shoots. It is superficially similar to the shrubby Desmanthodium perfoliatum, the latter readily distinguished by its markedly perfoliate leaves. Occasional plants of D. fruticosum may possess ternate leaves (e.g., Sundberg 2988 [NY]). volume 80(4):257-272 PHYTOLOGIA Apmil 1996 268 Figure 6. Distribution of Desmanthodium perfoliatum. Turner: . Revision of Desmanthodium 269 Desmanthodium fruticosum is closely related to D. guatemalense and the two might be reasonably combined. Both have shrubby habits and similar foliage, but the leaves of the latter are generally larger, thicker, and more nearly elliptic and less tapered at the base. Leaves on secondary shoots of both species tend to be smaller and narrower, making distinctions between these difficult. The two taxa are seemingly best distinguished by characters of the capitulescence, those of D. frulicosum possessing generally more numerous, smaller ultimate glomerules, the latter arranged in larger, more open, rounded cymose panicles. But, it must be admitted that occasional plants of D. fruticosum (e.g., Templeton 7643 [MICH], from Durango), were these collected in Guatemala, because of their large heads and foliage, would surely have been annotated as D. guatemalense. 5. DESMANTHODIUM TOMENTOSUM T.S. Brandegee, Univ. Calif. Publ. Bot. 6:73. 1914. TYPE: MEXICO. Chiapas: Cerro del Boquerén, Sep 1913, Purpus 6683 (HOLOTYPE: UC!). Desmanthodium congestum Arriagada & Stuessy, Brittonia 42:283. 1990. TYPE: MEXICO. Chiapas: San Vicente, 500 m, Aug 1938, E. Matuda 2508 (HOLOTYPE: GH; Isotypes: LL!,MICH!). Shrubs or subshrubs 1-3 m high; much resembling Desmanthodium guatemalense but the leaves thinner with somewhat larger blades which gradually taper onto the petioles, the latter 15-50 mm long; in addition the branches of the capitulescence are pubescent throughout (as opposed to pubescent in lines) with spreading tomentose hairs, as are the major veins beneath the blade. DISTRIBUTION (Figure 4) AND ECOLOGY: Known only from southwesternmost Chiapas and closely adjacent Guatemala in pine-oak forests, 1500- 2200 m; Aug-Nov. REPRESENTATIVE SPECIMENS: MEXICO. Chiapas: Mpio. de Uni6n, Faldas del Volcdn Tacana, 1500-1680 m, 18 Oct 1985, Villasefor Rios 864 (MEXU,TEX); SE side of Volcan Tacana above Talquian, 2200 m, 12 Nov 1972, Breedlove 29482 (MO). GUATEMALA: Prov. San Marcos: Finca Armenia, San Rafael pie de la Cuesta 10 Carrizal, past finca Africa, 1300-1600 m, 9-12 Aug 1980, Dwyer 15316 (MO). Amiagada & Stuessy (1990) thought their newly described Desmanthodium congestum to be sufficiently distinct so as to belong to a newly erected section Multiaggregata. They compared their taxon with both D. tomentosum and D. perfoliatum, noting its closer relationship with the former. Indeed, | am unable to distinguish D. congestum from D. tomentosum; their emphasis upon the more congested heads of the former is, in my opinion, illusionary, there being much vaniability and interpretational errors involved in ascertaining the number of heads involved in the ultimate aggregations of any given capitulescence, these varying from 4 to 24. _ Desmanthodium tomentosum is sufficiently close to D. guatemalense so that a case might be made for their treatment as but varieties of a single species. Indeed, the type 270 PHY TOLOGLA Apnil 1996 volume 80(4):257-272 of D. congestum itself, in pubescence, stands somewhere between these two taxa, although somewhat closer to D. tomentosum in leaf shape and texture. 6. DESMANTHODIUM GUATEMALENSE Hemsl., Biol. Centr. Amer. Bot. 2:142. t. 45. 1881. TYPE: GUATEMALA. Sacatepéquez: Volcan de Fuego, 6000 ft, (1860-1865), Salvin s.n. (HOLOTYPE: K). Desmanthodium hondurense A. Molina, Ceiba 11:70. 1965. TY PE: HONDURAS. Comayagua: Barranco Trincheras, 1200 m, 28 Dec 1952, Williams & Williams 18701 (HOLOTYPE: F). Stiffly erect to subscandent shrubs to 3 m high. Much resembling Desmanthodium fruticosum but the midstem leaves mostly thicker, more nearly elliptic, broadest at or near the middle, the petioles shorter (unwinged portion), and the heads generally larger, arranged in rather flat-topped or broadly rounded capitulescences, mostly broader than wide. DISTRIBUTION (Figure 4) AND ECOLOGY: Thickets along streams and slopes of volcanic cones, reportedly growing on volcanic ash in montane forests, 1200-3000 m; Jul-Nov. REPRESENTATIVE SPECIMENS: GUATEMALA. Baja Verapaz: King 3293 (TEX,UC). Chiquimula: Molina R. 26812 (US). San Marcos: Dwyer 15316 (US). Sacatepéquez: Croat 41985 (MO,US). Suchitepéquez: Skutch 1515 (LL). HONDURAS: Comayagua: Molina R. 31656 (MO). Itbuca: Rodriguez 81 (MO). Ocotepeque: Molina R. 30886 (MO). EL SALVADOR: Santa Ana: Croat 42367 (UC). Typical elements of this Central American species (as shown in Figure 4), possess large relatively thick leaves upon their primary stems, the blade tapering upon the petiole nearly to its base. Additionally, the heads are arranged on relatively short, thick branches which tend to form a stout, broad, somewhat flattened capitulescence. Nevertheless, secondary branches often possess much smaller leaves, the blades gradually tapering upon longer petioles. The latter, often populational, forms have been given the name Desmanthodium hondurense. Such forms are vegetatively very similar to D. fruticosum, but the two taxa are readily distinguished by their capitulescence, as noted in the above. Leaf variation on pnmary and secondary branches of D. guatemalense is neatly illustrated by M. Pahl in his delineation of the species for the Flora of Guatemala. 7. DESMANTHODIUM BLEPHARODON S.F. Blake, J. Wash. Acad. Sci. 14:454. 1924. TYPE: VENEZUELA. Tnyillo: “Between La Puerta and Timotes”, 2000 m, 16 Sep 1922, Alfredo Jan 1143 (HOLOTYPE: US). Much resembling Desmanthodium guatemalense but the leaves reportedly thicker, ovate, with 3-5 principal nerves, the blades broader near the base and pubescent along the lower margins; additionally the heads appear to be arranged in a rather evenly- Turner: _ Revision of Desmanthodium 271 fasciculate manner, resembling the capitulescence of species belonging to the remotely related genus Stevia. DISTRIBUTION (Figure 5) AND ECOLOGY: Known from only a few collections in Venezuela where it is “endémica de los paramos”, according to Anisteguieta (1964); Sep. According to its author, this taxon is nearest Desmanthodium guatemalense, the latter having “glaucescent branches and thin-membranous, more or less rhombic- ovate, subsessile leaves which are not ciliate at the base.” | Desmanthodium blepharodon is known to me only by the description and by an illustration in Ansteguieta (1964). It does appear very close to D. guatemalense but is maintained here because of its geographical isolation and because of its thicker, basally ciliate leaves. EXCLUDED SPECIES Desmanthodium trianae Hieron. = Clibadium trianae (Hieron.) S.F. Blake, Contr. Gray Herb., n. ser. 52:6. 1917. ACKNOWLEDGMENTS I am grateful to the various herbaria for the loan of specimens (GH, LL, MEXU, MICH, NY, UC, TEX) upon which the maps are based. Maria Thompson provided the illustration of Desmanthodium hintoniorum. Tom Wendt and Justin Williams reviewed the paper. LITERATURE CITED Anisteguieta, L. 1964. Desmanthodium, in Flora de Venezuela 10:383-384. Fay, J.J. 1974. In, IOPB Chromosome number reports XLV. Taxon 23:619-624. Kiel, D.J., et al. 1988. Chromosome studies in Asteraceae from the United States, Mexico, The West Indies and South America. Amer. J. Bot. 75:652-668. ee ey a rd ae. eley: 1977. Chromosome counts of Compositae from Mexico and the United States. Amer. J. Bot. 64:791-798. McVaugh, R. 1984. Desmanthodium, in Fl. Novo-Galiciana 12:294-296. University of Michigan Press, Ann Arbor, Michigan. Ralston, B.G., et al. 1989. Documented plant chromosome numbers 1989: Chromosome numbers in Mexican Asteraceae with special reference to the tribe Tageteae. Sida 13:359-368. Robinson, H. 1981. A revision of the tribal and subtribal limits of the Heliantheae (Asteraceae). Smithsonian Contr. Bot. 51:1-102. 272 PHY TOLOGIA Apni 1996 volume 80(4):257-272 Rydberg, P.A. 1915. Flaveria, in N. Amer. Fl. 34:142-146. Stuessy, T.F. & J.E. Armmagada. 1993. Chromosome counts in Clibadium (Compositae, Heliantheae) from Latin America. Brittonia 45:172-176. Sundberg, S., et al. 1986. Chromosome counts of Latin Amencan Compositae. Amer. J. Bot. 73:33-38. Tumer, B.L. 1996. The Comps of Mexico: Vol. 6. Phytologia Memoirs 10:1-93. Phytologia, Huntsville, Texas. Phytologia (April 1996) 80(4):273-275S. RECOGNITION OF TRIBES CAPSICEAE AND PHYSALEAE, SUBFAMILY SOLANOIDEAE, SOLANACEAE William G. D’Arcy & John E. Averett Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166 U.S.A. Department of Biology, Georgia Southern University, Statesboro, Georgia 30460 U.S.A. ABSTRACT Two new tribes in Solanaceae are described to accommodate new systematic information about the family. Capsiceae is centered on Capsicum and related genera, while Physaleae is centered on Physalis and related genera KEY WORDS: Solanaceae, Capsiceae, Physaleae, Solanoideae, taxonomy The genera Capsicum L. and Physalis L., family Solanaceae, subfamily Solanoideae, have traditionally been placed in tribe Solaneae (Wettstein 1891). They are distanced from Solanum L. (type species of the family and of the Solaneae) and most other members of the tribe by distinctive calyces and other features, and the following recognizes their distinctiveness at the tribal level. Capsiceae D’Arcy, tribus nov. Type genus: Capsicum L., Gen. Pl., ed. 5:86. 1754. Herbae perennes vel frutices. Flores campanulati vel rotati. Calyces truncati, interdum dentes subapicales ferenti. Fructus baccati vel drupacei. The calyx in Capsicum lacks the terminal lobes found in other Solanaceae but is apically truncate, sometimes with subapical enations that may resemble terminal lobes. The calyx is hardly accrescent in fruit. In his revision of Lycianthes, Bitter (1920) suggested a close relationship between Lycianthes and Capsicum, and subsequent morphological studies of the diagnostic calyces (D’Arcy 1986; Bernardello & Hunziker 1987) support this, arguing that these two genera form a core group of the new tribe Capsiceae. D’Arcy (1991) suggested that the following genera have calyx features at least superficially similar to Capsicum and may also belong to this new PA fo) 274 PHYTOLOGIA Apni 1996 volume 80(4):273-273 tribe: Acnistus, Aureliana, Dunalia, lochroma, Lycianthes, Saracha, Tubocapsicum, Vassobia, and Witheringia. Physaleae D’Arcy, fribus nov. Type genus: Physalis L., Gen. Pl., ed. 5: 85. 1754. Herbae perennes vel frutices. Flores campanulati vel rotati. Calyces dentes terminales ferenti in statu fructu accrescenti. Fructus baccati vel drupacei. The calyx in Physalis has terminal lobes and is accrescent in fruit, surrounding the berry. Anthers are longitudinally dehiscent, ovaries have nectaries, and plants never have prickles. In Solanum, calyces are only exceptionally accrescent (S. sisymbrifolium Lam., S. toliarea D’Arcy & Rakotozafy), anthers are poricidal, ovaries lack nectaries, and plants are often prickly, a suite of characters indicating more than a generic taxonomic distance. , Accrescent calyces occur in some other genera of Solanoideae, (e.g., Witheringia folliculoides J.L. Gentry & D’Arcy and Nicandra physaloides [L.] Gaertner). This suggests that accrescent calyces may be plesiomorphic in subfamily Solanoideae, appearing as a conservative condition in a few groups, or contranly that accrescent calyces have risen independently in each of these lineages. In either case, accrescent calyces remain a single useful character to suggest the inclusion of Physalis and other genera in the new tribe, Physaleae. The following genera, which have similar accrescent calyces, are sometimes referred to as the ‘physaloid group’ and were suggested by Averett (1977, 1979), D’Arcy (1991) or Axelius (1996) to be closely allied to Physalis. They may also belong to this new tribe: Archiphysalis, Brachistus, Chamaesaracha, Deprea, Jaltomata, Leucophysalis, Margaranthus, Melilissia, Quincula, Physaliastrum, and Withania. LITERATURE CITED Averett, J.E. 1977. Taxonomic notes and new combinations in Leucophysalis (Solanaceae). Ann. Missour Bot. Gard. 64:141-142. Averett, J.E. 1979. The North American Genera of the Solaneae in J.E. Hawkes (ed.). The Biology and Taxonomy of the Solanaceae., Academic Press, pp. 493- 512. Axelius, B. 1996. The phylogenetic relationships of the physaloid genera (Solanaceae) based on morphological data. Amer. J. Bot. 83:118-124. Bernardello, L.M. & A.T. Hunziker. 1987. Estudios sobre Solanaceae XXXIII. género Lycianthes en La Argentina. Darwiniana 31:17-34. Bitter, G. 1920. Die Gattung Lycianthes. Abh. Naturwiss. Vereine Bremen 24:292- 520. D’Arcy, W.G. 1986. The calyx in Lycianthes and some other genera. Ann. Missouri Bot. Gard. 73:117-127. D’Arcy & Averett: _ New tribes in Solanaceae 275 D’Arcy, W.G. 1991. The Solanaceae since Birmingham, 1976 with a review of its biogeography. pp. 75-137 in J.G. Hawkes, R. Lester, M. Nee, & N. Estrada (eds.) Solanaceae 3: Taxonomy-Chemistry-Evolution. Royal Botanical Gardens, Richmond, United Kingdom. Wettstein, R. von. 1891. Solanaceae. vol. 4(3b):1-38. in A. Engler & K. Prantl. Die Natiirlichen Pflanzenfamilien. Leipzig, Germany. Phytologia (April 1996) 80(4):276-279. A NEW SPECIES OF ROLDANA (SENECIONEAE) FROM OAXACA, MEXICO B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A new species, Roldana calzadana B.L. Turner, is described and illustrated from Oaxaca, México. It is closely related to the recently described Senecio galicianus McVaugh var. manantlanensis Kowal, which is endemic to the Sierra Manantlan of Jalisco. The latter is elevated to specific rank and treated as belonging to the genus Roldana, thus necessitating the following name change: Roldana manantlanensis (Kowal) B.L. Turner, stat. & comb. nov. In addition, the following new combinations within Roldana are proposed: R. gonzalezae (B.L. Tumer) B.L. Turner, comb. nov.; R. neogibsonii (B.L. Turner) B.L. Turner, comb. nov.; and R. sundbergii (B.L. Turner) B.L. Turner, comb nov. KEY WORDS: Asteraceae, Senecioneae, México, Oaxaca, Senecio, Roldana, systematics Studies on the Asteraceae of México have necessitated the following descriptions and name changes. ROLDANA CALZADANA B.L. Turner, spec. nov. Figure 1. TYPE: MEXICO. Oaxaca: Mpio. San Martin Peras, carretera Coicoyan de las Flores - Santiago Juxtlahuaca (17° 17’ N x 98° 11’ W), “200 m de la deviacion a San Martin Peras”, pine-oak woodland, ca. 2535 m, 16 Feb 1995, J.J. Calzada 19738 (HOLOTYPE: TEX!; Isotype: MEXU). Similis Roldanae manantlanensis (Kowal) B.L. Turer sed habens folis graciliora, lobis deltatioribus et irrgulaniter dentatis. 276 277 New Roldana from Oaxaca Tumer. Figure 1. Roldana calzadana, from holotype. 278 PHYTOLOGIA Apnil 1996 volume 80(4):276-279 Suffruticose herbs to 2 m high. Stems (upper) tawny-puberulous, pithy. Leaves (larger) 30-35 cm long, 16-20 cm wide; petioles 12-15 cm long; blades broadly ovate- elliptic in outline, cordate basally, 7-9 palmately nervate from the base, both surfaces glabrous, except along the major veins, the lateral margins bearing 5-6 deltoid lobes, each of the latter irregularly serrate. Heads arranged in rather flat-topped congested cymes ca. 6 cm high, 9 cm wide, the ultimate peduncles sparsely tomentose, 4-10 mm long. Involucres cylindro-campanulate, 5-6 mm high. Involucral bracts ca. 8, their apices greenish and broadly deltoid. Ray florets 3 per head; ligules yellow, 4-6 mm long, 1.5-2.5 mm wide; tubes puberulent. Disk florets 5 per head; corollas yellow, 7- 8 mm long, sparsely pubescent. Achenes (immature) columnar, ca. 2 mm long, glabrous. Pappus of numerous readily deciduous white bristles 5-6 mm long. This taxon is closely related to the recently described Senecio galicianus McVaugh var. manantlanensis Kowal, from Sierra Manantlan, Jalisco (cf. below). They possess similar habits, leaves and involucres, and both have similar florets with pubescent corollas. Roldana calzadana differs in having leaves with more broadly deltoid marginal lobes, each irregularly serrate, and heads arranged in rather congested flat-topped cymes which are over-topped by the upper foliage. In my preliminary treatment of the Roldana complex for the comps of México (cf. Turner 1996), Barkley (in our collaborative work on Senecio, s.|.) prevailed upon me to treat Roldana within the broad bounds of his concept of Senecio (Barkley 1985); more recently he has come full circle and would treat Roldana (among many other segregates) as generically distinct. This has necessitated the following name changes: ROLDANA GONZALEZAE (B.L. Turner) B.L. Turner, comb. nov. Based upon Senecio gonzalezae B.L. Turner, Phytologia 57:377. 1985. ROLDANA NEOGIBSONII (B.L. Turner) B.L. Turner, comb. nov. Based upon Senecio neogibsonii B.L. Turner, Brittonia 37:119. 1985. ROLDANA SUNDBERGII (B.L. Turner) B.L. Turner, comb. nov. Based upon Senecio sundbergii B.L. Turner, Brittonia 37:117. 1985. ROLDANA MANANTLANENSIS (Kowal) B.L. Turner, comb. & stat. nov. Based upon Senecio galicianus McVaugh var. manantilanensis Kowal, Bnittonia 43:109. 1991. Kowal (1991) has provided a tedious, detailed, wonderfully elaborated upon, account documenting the biological reality of this taxon. He treats it as a variety of Senecio galicianus McVaugh but I think it deserving of specific rank and will recognize it as such in my treatment of Roldana for the Comps of Mexico (vol, 7, in prep). Jeffrey (1992) transferred several other of my roldanoid species of Senecio into Roldana: these include Senecio carlomansonii B.L. Tumer & T. Barkley; S. Tumer.: New Roldana from Oaxaca 279 gesneriifolius B.L. Turner 1987 (non S. gesneriifolius Cuatrec. 1950), but graciously given the new name R. gesneriifolius C. Jeffrey; S. grimesii B.L. Turner; S. marquesii B.L. Turner; S. metapecus B.L. Turner; and S. nesomiorum B.L. Turner. ACKNOWLEDGMENTS I am grateful to WIS for the loan of relevant material relating to the present paper. Gayle Turner provided the Latin diagnosis, and she and Justin Williams reviewed the paper. LITERATURE CITED Barkley, T. 1985. Infrageneric groups in Senecio s.l. and Cacalia s.|. (Asteraceae: Senecioneae) in Mexico and Central America. Brittonia 37:211-218. Jeffrey, C. 1992. Notes on Compositae VI. Kew Bull. 47:49-109. Kowal, R.K. 1991. A new vanety of Senecio (Asteraceae: Senecioneae) from the Sierra de Manantlan, Jalisco, Mexico, with notes on the S. roldana complex. Brittonia 43: 102-115. Turner, B.L. 1996. The Comps of Mexico--Tageteae and Anthemideae. 6:1-93. Phytologia Memoirs 10, Phytologia, Huntsville, Texas. Phytologia (Apml 1996) 80(4):280-283. COMMENTS ON THE DISTRIBUTION OF BOTRYCHIUM LUNARIOIDES (OPHIOGLOSSACEAE) IN TEXAS W.C. Holmes, T.L. Morgan, J.R. Stevens, & R.D. Gooch Department of Biology, Baylor University, Waco, Texas 76798-7388 U.S.A. & J.R. Singhurst Wildlife Division, Texas Parks and Wildlife Department, Mexia, Texas 76667 tLS.A. ABSTRACT Botrychium lunarioides (Michx.) Sw. (Ophioglossaceae) is now known to be widespread and abundant throughout the eastern portion of Texas. KEY WORDS: Ophioglossaceae, Botrychium, Texas, biogeography Until recently, Botrychium lunarioides (Michx.) Sw. (Ophioglossaceae) was considered to be a species of the coastal plain of the southeastern United States whose westernmost distribution was known to extend to extreme east Texas (Thomas 1979; Thomas, et al. 1981; Wagner & Wagner 1993). In 1996, Do, ef al. reported ten additional county records in the central portion of the Post Oak Savannah of Texas, thereby extending the known distribution of the species up to 273 km to the west. Additional field studies during 1996 have yielded nineteen new county records for the species in Texas (Figure 1). These new reports are primarily from the Post Oak Savannah, Pineywoods, and Blackland Prairies of northeast Texas and from the southern portion of the Post Oak Savannah. The most notable occurrence of the species is at Lake Bastrop State Recreation Area, Bastrop County, about 45 km ESE of Austin, which extends both the western and southern known limits of the species. The present distribution confirms that the plant is much more widespread and abundant within the state than previously known and that it may be expected to occur in nearly all counties of the Pineywoods and the Post Oak Savannah. Although the habitat of the very southern portion of the Post Oak Savannah (Caldwell, Gonzalez, Guadalupe, and Wilson counties), seems to be favorable for the species, an attempt to locate it Holmes et al.: Distribution of Botrychium lunarioides in Texas Figure 1. Documented distribution of Botrychium lunarioides in Texas. 281 282 PHYTOLOGIA April 1996 volume 80(4):280-283 there was unsuccessful. This failure may be related to the extreme drought in that area in the late winter and early spring of 1996, which has been mentioned by Thomas, ef al. (1981) as a factor that may cause the species to remain dormant. Specimens Examined: Texas: Anderson Co.: Cedar Creek Cemetery, between Long Lake and Elkhart on Texas Hwy. 294, 135 m, 14 Mar 1995, Holmes 7602 (BAYLU). Bastrop Co.: Lake Bastrop State Recreation Area, ca. 0.6 mile S of jet. of FM 1441 and N Shore Park Road, lawn NE of boat ramp, 13 Mar 1996, Singhurst 4806 (BAYLU). Bowie Co.: ca. 2.6 miles E of jct. of U.S. Hwy. 67 and FM 98 at Simms, Old Martin Cemetery, 6 Mar 1996, Singhurst 4805 (BAYLU). Camp Co.: Rose Hill Cemetery, Tex. Hwy. 11 at jct. with U.S. Hwy. 271, 5 Apr 1996, Stevens 203 & Gooch (BAYLU). Cass Co.: Smyrna Cemetery on east side of Tex. Hwy. 77, ca. 4 miles NW of the Louisiana State line NW of Rodessa, LA and SW of Atlanta, TX, 31 Mar 1988, Thomas 103695, Dorris, & Slaughter (NLU). Franklin Co.: Hogansport Cemetery on County Road NW 1028 ca. 0.1 mile S of FR 71, 5 Apr 1996, Stevens 200 & Gooch (BAYLU). Freestone Co.: Fairfield Lake State Recreation Area, 6 miles NE of Fairfield, Chancellor Cemetery, 22 Feb 1995, Do 324 (BAYLU); New Hope Cemetery, 18 Mar 1995, Singhurst 3029 (BAYLU). Falls Co.: Williams Cemetery, ca. 5 miles S of Kosse on Texas Hwy. 14, 17 Mar 1995, Singhurst 3004 (BAYLU). Henderson Co.: Ash Cemetery, just SE of Murchison on Tex. Hwy. 31, 17 Mar 1995, Holmes 7610 (BAYLU). Hopkins Co.: Harmony Cemetery on County Road 2397 ca. 0.5 miles NW of County Road 2403, 5 Apr 1996, Stevens 206 & Gooch (BAYLU). Houston Co.: ca. 0.2 mile N of jet. Tex. Hwy. 21 and park road, ca. 5 miles NW of L.R. Price Log Cabin; 29 Jan 1996, Singhurst 4734 (BAYLU). Hunt Co.: Donelton Cemetery*on County Road 3219 ca. 0.8 mile NW of FR 1567, 6 Apr 1996, Stevens 208 & Gooch (BAYLU). Kaufman Co.: New Salem Cemetery on County Road 315 ca. 1.5 miles N of Interstate Hwy. 20, 6 Apr 1996, Stevens 210 & Gooch (BAYLU). Lee Co.: Tanglewood Cemetery, ca. 5.2 miles N of jct. of U.S. Hwy. 77 and FM 696, ca. 0.2 mile W and N on unnamed road ca. 0.4 mile, 31 Jan 1996, Singhurst 4773 (BAYLU). Leon Co.: FR 3, 1.5 miles S of jct. with U.S. Hwy. 79 at Winn Cemetery, 110 m, 3 Mar 1995, Gooch 63, Stevens, & Holmes (BAYLU). Limestone Co.: McKenzie Cemetery, 12 Mar 1995, Singhurst 3024 (BAYLU); Cobb Cemetery, 12 Mar 1995, Singhurst 3024 (BAYLU); Cobb Cemetery, 12 Mar 1995, Singhurst 3025 (BAYLU); Ferguson Cemetery, off FR 937, ca. 2 miles NW of the Robertson Co. line, 12 Mar 1995, Singhurst 3020 (BAYLU). Milam Co.: Old Providence Cemetery, 20 Mar 1995, Singhurst 3201 (ASTC). Morris Co.: Daingerfield State Park, ca. 2.1 miles S of jet. of Tex. Hwy. 49 and park road, lawn area ca. 80 m NW of activity center, 7 Mar 1996, Singhurst 480] (BAYLU). Navarro Co: Midway Cemetery, ca. 7 miles NE of Streetman, 18 Mar 1995, Singhurst 3028 (BAYLU). Rains Co.: Prospect Cemetery on County Road 1230 ca. 0.5 mile S of U.S. Hwy. 69, 6 Apr 1996, Stevens 209 & Gooch (BAYLU). Red River Co.: McCrary Cemetery on FR 196 ca. 2 miles W of Tex. Hwy. 37, 5 Apr 1996, Stevens 198 & Gooch (BAYLU). Robertson Co.: FR 979 at jct. with FR 2096 at Bald Prairie Cemetery, 98 m, 3 Mar 1995, Stevens 82, Gooch, & Holmes (BAYLU); Seale Round Prairie Cemetery, off FR 937, ca. 2 miles SE of the Limestone Co. line, 12 Mar 1995, Singhurst 3022 (BAYLU). Rusk Co.: Martin Creek State Park, ca. 1.3 miles SW of jct. of FM 1716 and park road, lawn next to cabin no. 1, 5 Mar 1996, Singhurst 4804 (BAYLU). San Augustine Co.: Liberty Hill Baptist Church Cemetery, 2.3 miles N of Tex. Hwy. 21 by Tex. Hwy. 147, 16 Feb 1972, Thomas 27495 (NLU). Smith Co.: Tyler State Park, 0.6 mile W Holmes et al.: Distribution of Botrychium lunarioides in Texas ye of jct. of FR 14 and park road, lawn S of registration office, 4 Mar 1996, Singhurst 4803 (BAYLU). Titus Co.: Winfield Cemetery on Interstate Hwy. 30 ca. 200 m E of Spur 185, 5 Apr 1996, Stevens 20] & Gooch(BAYLU). Upshur Co.: Cemetery on U.S. Hwy. 271 ca. 1 mile S of Camp Co. line, 5 Apr 1996, Stevens 204 & Gooch (BAYLU). Van Zandt Co.: Purtis Creek State Recreation Area, at jct. of FM 316 and Gosham Road, 4 May 1995, Singhurst 3261 (BAYLU). Walker Co.: Huntsville State Park, jct. of Interstate Hwy. 45 and park road, ca. 1.9 miles SW of park road, N side of park road in lawn E of education center, 28 Feb 1996, Singhurst 4800 (BAYLU). Waller Co.: Macedonia Cemetery, E off of Macedonia School Road, N of Magnolia Road and S of Threemile Creek near the intersection of Harris, Walker and Montgomery cos., 19 Mar 1992, Brown 15832 (BAYLU (photo),SBSC). Wood Co.: Perryville Cemetery on FR 852 ca. 0.5 mile S of FR 1647, 5 Apr 1996, Stevens 205 & Gooch (BAYLU). ACKNOWLEDGMENTS We wish to thank Larry E. Brown of SBSC for the loan of a specimen from Waller County, the Texas Parks and Wildlife Department for access to public lands under their jurisdiction, and the Beta Tau Chapter of Beta Beta Beta of Baylor University for partial financial support provided through a Bob Gardner Memonal research Grant to Stevens and Gooch. LITERATURE CITED Do, L.H., R.D. Gooch, J.R. Stevens, W.C. Holmes, & J.R. Singhurst. 1996. New county records of Botrychium lunarioides in Texas. Amer. Fern J. 86:28-31. Thomas, R.D. 1979. First record of Botrychium lunarioides and Ophioglossum nudicale var. tenerum (Ophioglossaceae) from Texas. The Southw. Naturalist 24:395-396. Thomas, R.D., T. Briley, & N. Carroll. 1981. Additional collections of Botrychium lunarioides from Texas and Oklahoma and comments on its dormancy. Phytologia 48:276-278. Wagner, W.H. & F.S. Wagner. 1993. Jn Flora of North America Editorial Committee, eds. Flora of North America, vol. 2, Pteridophytes and Gymnosperms. Oxford Press, New York, New York (Pp. 85-106). Phytologia (April 1996) 80(4):284. NEW COMBINATIONS FOR THE FLORA OF THE CENTRAL EASTERN UNITED STATES Clyde F. Reed 1222 Main St., Darlington, Maryland, 21034-1416 U.S.A. ABSTRACT New combinations are made for Panicum acuminatum var. wrightianum and P. sphaerocarpon var. isophyllum. KEY WORDS: Poaceae, Panicum, Easter United States, nomenclature The following new combinations are necessary for the completion of a manuscript for the Flora of the Central Eastern United States. |. Panicum acuminatum Swartz var. wrightianum (Scribn.) Reed, comb. nov. BASIONYM: Panicum wrightianum Scribn., U.S.D.A., Div. Agrost. Bull. 11:44, f. 4. 1898. 2. Panicum sphaerocarpon Elliott var. isophyllum (Scribn.) Reed, comb. nov. BASIONYM: Panicum microcarpon Muhl. var. isophyllum Scribn., Bull. Tenn. Agric. Exp. Sta. 7:51. f. 54. 1894. Phytologia (April 1996) 80(4):285-287. A NEW SPECIES OF GALIUM (RUBIACEAE) FROM NORTHEASTERN MEXICO B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT Galium hintoniorum B.L. Tumer, spec. nov., is described and illustrated. It is a prostrate rhizomatous herb with numerous relatively small leaves and is known only from southernmost Tamaulipas, México, where it occurs in oak woodlands between 1300 and 1900 meters. KEY WORDS: Rubiaceae, Galium, México, Tamaulipas, systematics Routine identification of plants from the Sierra Madre Onental of northeastern Mexico has revealed the following novelty. GALIUM HINTONIORUM B.L. Turmer, spec. nov. Figure 1. TYPE: MEXICO. Tamaulipas: Mpio. Guemes, Los Pedros, “Grass savannah in oak woods,” 1355 m, 10 Nov 1994, Hinton, et al. 25127 (HOLOTYPE: TEX!). Simile G. microphyllo A. Gray sed foliis plerumque parvioribus, nigrescentibus in sicco (vice viridium), apicibus foliorum tantum acutatis (vice apicum apiculatorum), et fructificationibus pubescentibus (vice glabrarum) cum pilis parvis arcuatisque. Prostrate rhizomatous herbs 10 cm high or less. Stems moderately white-pilose with spreading hairs to densely ciliate with upcurved arcuate hairs, the vestiture 0. 1- 0.3 mm high. Leaves 4 to a node throughout, numerous and much overlapping; petioles ca. 0.25 mm long; blades ovate-elliptic to narrowly oblanceolate, mostly 3-9 mm long, 1-2 mm wide, uninervate, glabrous or nearly so, shiny, the margins entire and thickened, the apices merely acute. Flowers few, mostly axillary and shortly pedicellate but some of them seemingly terminal and sessile. Flowers rotate campanulate, the petals 2-3 mm long, glabrous. Fruiting bodies ca. 1.2 mm long, moderately and evenly ornate with arcuate upcurved hairs. 285 286 PHYTOLOGIA Apnil 1996 volume 80(4):285-287 Wy, MT gl a a4 ca fi PPh” be ES tn é MA) Cape, AG ) hee aay - > 2 aes 3 TAS 4 WE, 4 Dy 24) i iF a XD \} DEAT i SA A n) Ly W\4q BRU fb YT y, \ 7 ¥ $3: IN CE SOVA RIDA i FFM Af 71,4 (I ly a Se) 7 Ke rer =. 7 oA ; 2 ' a Ve ZV a ee | eS AY ZA) \ q ff 7 Le yy, Ag ) Ze YVR <7 } — Wa hy 1 Sb = NG BN\ ar4 ZB y we Figure 1. Galium hintoniorum, from holotype; upper right, portion of midstem showing leaves; lower right, fruiting body. Tumer: New Galium from México 287 ADDITIONAL SPECIMENS EXAMINED: MEXICO. Tamaulipas: Mpio. Hidalgo, Los Caballos, 1705 m, 21 Sep 1994, Hinton, et al. 24804 (TEX); road from Sta. Engracia toward Dulces Nombres, N.L., 0.3 mi W of Paraje de Los Caballos (22° 58’ 39” N x 99° 29’ 31” W), 1840 m, 21 Sep 1994, Nesom 7460 with Jaime Hinton & M. Mayfield) (TEX). Galium hintoniorum much resembles G. microphyllum but it apparently lacks the 4-bracted flowers of that taxon, and the fruits are ornamented with recurved hairs, as illustrated in Figure 1. Additionally, the foliage dries black and is more densely packed, the leaves lacking whip-like acuminations at their apices as occur in G. microphyllum. In short, the species appears to stand somewhere between the Relubium taxa (Galium microphyllum, et al.) and the more typical elements of Galium, seemingly Vitiating the characters upon which these two genera are founded. The species is named for the Hinton family, all of the collections having been made by or with Jaime Hinton, the extant patriarch of that remarkable clan. ACKNOWLEDGMENTS I am grateful to Gayle Tumer for the Latin diagnosis, and to her and Ted Delevoryas for reviewing the manuscript. Maria Thompson provided the illustration. Phytologia (Apml 1996) 80(4):288-290. NOMENCLATURAL COMBINATIONS IN CYPERUS (CYPERACEAE) Stanley D. Jones!, Joseph K. Wipff 1, & Richard Carter2 1BRCH Herbarium, Botanical Research Center, P.O. Box 6717, Bryan, Texas 77805-6717 U.S.A. e-mail: sdjones@bihs.net; wipff@bihs.net 2VSC Herbarium, Biology Department, Valdosta State College, Valdosta, Georgia 31698 U.S.A. e-mail: RCarter@ grits. valdosta. peachnet.edu ABSTRACT The following nomenclatural changes in Cyperaceae are proposed: Cyperus macrocephalus F. Liebmann var. eggersii (J. Bockeler) comb. nov.; Cyperus odoratus C. Linnaeus var. engelmannii (E. von Steudel) comb. et stat. nov.; and Cyperus odoratus C. Linnaeus var. squarrosus (J. Bockeler) comb. nov. KEY WORDS: Cyperaceae, Cyperus, nomenclature, Cyperus eggersii, Cyperus macrocephalus, Cyperus odoratus var. engelmannii, Cyperus odoratus var. squarrosus, subgenus Diclidium Tucker (1994) treated Cyperus eggersii J. Bockeler as a synonym of C. odoratus C. Linnaeus, subgenus Diclidium (Nees von Esenbeck) C.B. Clarke [Sy = subgenus Torulinium (N. Desvaux) G. Kiikenthal]. Adams (1991, 1994) recognized subgenus Torulinium at the rank of genus and placed Cyperus eggersii as a variety of Torulinium macrocephalum (F. Liebmann) C.B. Clarke. We agree with Tucker’s placement of subgenus Diclidium (Sy = Torulinium) but agree with Adams that Cyperus eggersii should be recognized as a variety of C. macrocephalus F. Liebmann. However, since Adams recognized subgenus Torulinium as a distinct genus, a new combination in Cyperus becomes necessary. We have examined herbarium specimens and field populations of all three taxa from Texas south to Belize. We have found but a single population appearing as a putative hybrid between C. eggersii and C. ferruginescens J. Bockeler and have had no problems differentiating one taxon from another. This is not to say that somewhere, mixed populations and intermediates may exist. However, we have done extensive field and herbarium work and do not see the justification for treating these three taxa as a single entity. 288 Jones, et al.: New combinations in Cyperus 289 Cyperus macrocephalus F. Liebmann var. eggersii (J. Bockeler) S.D. Jones, J. Wipff, & R. Carter, comb. nov. BASIONYM: Cyperus eggersii J. Bockeler, Beitrage zur Kenntniss der Cyperaceen 1:53. 1888. Torulinium eggersii (J. Bockeler) C.B. Clarke in I. Urban (Editor), Symbolae Antillanae Seu Fundamenta Florae Indiae Occidentalis 2:56. 1900-1901. Torulinium macrocephalum (F. Liebmann) T. Koyama var. eggersii (J. Bockeler) C.D. Adams, Annals of the Missouri Botanical Garden 78:254. Toot, TYPUS: REPUBLICA DOMINICANA. Santo Domingo: near Batey on Rio Yasica, 23 Jun 1887, Eggers 2627 (HOLOTYPE: B). Many North American authors since Fernald (1950) have treated Cyperus engelmannii E. von Steudel and C. ferruginescens J. Bockeler as synonyms of C. odoratus, usually without explanation. Two notable exceptions are Braun 1967 and Voss 1972. Most authors refer to C. odoratus as a polymorphic species commenting that C. odoratus is the most variable Cyperus in their area. We find these three taxa closely related and mostly sympatric, but discrete. Although some intermediates exist, they are relatively few. Considering their distinct morphologies, we believe that varietal rank under C. odoratus is warranted for C. engelmannii and C. ferruginescens. The following combinations are made: Cyperus odoratus C. Linnaeus var. engelmannii (E. von Steudel) R. Carter, S.D. Jones, & J. Wipff, comb. et stat. nov. BASIONYM: Cyperus engelmannii E. von Steudel, Synopsis Plantarum Cyperacearum 2:47. 1854. Cyperus ferax L.C. Richard subsp. engelmannii (E. von Steudel) G. Kiikenthal, Pflanzenreich 4(20):620. 1936. TYPUS: UNITED STATES. Illinois: Cahokia, Sep 1845, G. Engelmann s.n. (HOLOTYPE: P?; Isotype: GH). Cyperus odoratus C. Linnaeus var. squarrosus (N. Britton) S.D. Jones, J. Wipff, & R. Carter, comb. nov. BASIONYM: Cyperus speciosus M.H. Vahl var. squarrosus N. Britton, Memoirs of the Torrey Botanical Club 13:214. 1886. Cyperus ferruginescens J. Bockeler, Linnaea. 36:396-397. 1869-70. Cyperus speciosus M.H. Vahl var. ferruginescens (J. Bockeler) N. Britton, Memoirs of the Torrey Botanical Club 5:61. 1894. TYPUS: UNITED STATES. Missouri: St. Louis. No specimen cited in protologue. The following key is modified from O'Neill (1940) and will differentiate the species of Cyperus subgenus Diclidium in Texas. KEY TO CYPERUS SUBGENUS DICLIDIUM IN TEXAS 1. Inflorescence consisting of a single aggregated subglobose head, the spikes sessile; in Texas, restricted to the Lower Rio Grande Valley and sparingly upward along the coast to the Coastal Bend........... C. macrocephalus var. macrocephalus 1. Inflorescence consisting of several peduncled spikes. 290 PHYTOLOGIA Apmil 1996 volume 80(4):288-290 2. Spikelets appressed to ascending, densely crowded; in Texas, restricted to the Lower Rio Grande Valley. ....................068. C. macrocephalus var. eggersii 2. Spikelets divaricate, scattered along the rachis of the spike; in Texas, widely distributed. 3. Scales near the middle of the spikelet (2.7-)2.8-3.2 mm long; rachilla wings reaching or covering the shoulders of the achene; achenes (1.2-)1.3-1.5 mm long, (0.5-)0.6-0.7 mm wide; spikelets RNIN ON is ached Sic nals Sareea enn one C. odoratus var. odoratus 3. Scales near the middle of the spikelet (2.0-)2.3-2.5(-2.6) mm long, rachilla wings rarely reaching and never covering the shoulders of the achene; achenes 0.8-1.0{-1.1) mm long, (0.3-)0.4-0.5 mm wide; spikelets reddish. 4. Tip of scale reaching only to base of the scale next above and on the same side of the rachis....................066 C. odoratus var. engelmannii 4. Tip of scale conspicuously reaching over the base of the scale next above on the same side of the rachis. ........ C. odoratus var. squarrosus ACKNOWLEDGMENTS We appreciate Paul A. Fryxell (TEX) and Gretchen D. Jones (USDA-ARS) for their manuscript review. We thank Dan Nicolson (Smithsonian) for his constructive comments on selected nomenclature. LITERATURE CITED Adams, C.D. 1991. New combinations and a new variety in the Cyperaceae of Mesoamerica. Annals of the Missouri Botanical Garden. 78(1):254. Adams, C.D. 1994. Torulinium. In. Flora Mesoamericana. Vol. 6. Alismataceae a Cyperaceae. Universidad Nacional Aut6noma de México; Instituto de Biologia, México City, México. Braun, E.L. 1967. The Monocotyledoneae: Cat-tails to Orchids. The Ohio University Press, Cincinnati, Ohio. Fernald, M.L. 1950. Gray’s Manual of Botany. 8th ed. Dioscorides Press, Portland, Oregon. O’Neill, H.T. 1940. Botany of the Maya Area: The sedges of the Yucatan Peninsula. Publication No. 19. Carnegie Institute of Washington, Washington, D.C. Tucker, G.C. 1994. Revision of the Mexican species of Cyperus (Cyperaceae). Systematic Botany Monographs 43: 1-213. Voss, E.G. 1972. Michigan Flora. Vol. 1. Monocotyledons. Cranbrook Institute, Bloomfield Hills, Michigan. Phytologia (Apmil 1996) 80(4):291-294. A NEW VARIETY OF COREOPSIS MUTICA (ASTERACEAE) FROM WESTERN OAXACA B.L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A new variety of Coreopsis mutica, C. mutica var. miahuatlana B.L. Turner, is described from Mpio. Miahuatlan, Oaxaca, México. It is a shrub or small tree 2.0-5.0 m high and relates closely to var. subvillosa. The distinctions between these two varieties are discussed. KEY WORDS: Asteraceae, Coreopsis, México, Oaxaca, systematics Crawford (1970) provided an excellent study of Coreopsis mutica A. DC. in which seven regional varieties were recognized. Crawford (1981) added an additional varietal element to the group. Turner (1992) reviewed the taxonomy of the complex, reducing some of Crawford’s taxa to synonymy, but added a newly described taxon, C. mutica var. guerreroana B.L. Tumer, maintaining seven varietal taxa, all of these keyed and mapped. I describe herein a new variety, Coreopsis mutica var. miahuatlana from west- central Oaxaca, where it is seemingly confined to Mpio. Miahuatlan. In my revised key to the group (Tumer 1992) the present taxon will key to var. carnosifolia D. Crawford, but differs from the latter in having much larger, less succulent leaves which dry dark (vs. semisucculent smaller leaves which dry pale green). In addition the capitulescence is broader and more numerous-headed. The closest relationship of var. miahuatlana appears to be with var. subvillosa D. Crawford; indeed, occasional specimens of the former take on characters of the latter (e.g., Hinton 26603, in pubescence; and Hinton 26735, in its lobed leaves). In my treatment of 1992 | included at least one collection of var. miahuatlana (Turner 80A-9 [TEX], cited below) in my concept of var. subvillosa, taking this at the tme to be an intermediate between var. subvillosa and var. carnosifolia. 1 would now call this collection an intergradant between the latter and var. miahuatlana. 291 292 PHYTOLOGIA April 1996 volume 80(4):291-294 Figure 1. Coreopsis mutica vat. miahuatlana (from holotype). Tumer: New variety of Coreopsis mutica from Oaxaca 293 COREOPSIS MUTICA A. DC. var. MIAHUATLANA B.L. Turner, var. nov. Figure 1. TYPE: MEXICO. Oaxaca: Mpio. Miahuatlén, Santo Domingo, 2240 m, IRF Rfo Magdalena, tree 5S m, common, 4 Aug 1996, Hinton, et al. 26724 (HOLOTYPE: TEX!). Similis Coreopsi muticae (Coreopsis mutica) A. DC. var. subvillosa D. Crawford, sed plantae majores sunt, foliis primariis majoribus, plerumque 10- 13 cm longis (vice 3-9 cm longis), denigratis ubi siccis, glabris aut paene glabris ubique, et foliis immaturis subvillosis. Shrub or small tree 2.0-5.0 m high. Stems and foliage glabrous throughout or nearly so, but juvenile leaves sometimes moderately to sparsely pubescent. Midstem (primary) leaves mostly 10-15 cm long, 2.5-4.5 cm wide, drying dark green or blackish, the margins serrulate. Heads numerous, arranged in broad rounded terminal corymbose panicles 5-10 cm high, 10-16 cm across. Primary peduncles 1-2 cm long, the ultimate peduncles mostly 1-3 cm long. Involucres narrowly campanulate, 7-9 mm high, 5-6 mm wide (pressed); outer bracts 4-5, oblanceolate, 3-nervate, mostly 4-7 mm long; inner bracts ca. 8, subscarious, free to base or nearly so. Ray florets mostly 5; ligules yellow, 1-2 cm long, 0.6-1.0 cm wide. Disk florets 18-25 (estimated); corollas yellow. Anther sacs purple. Achenes narrowly ellipsoidal, ca. 8 mm long, 2 mm wide. ADDITIONAL SPECIMENS EXAMINED: (all from Mpio. Méiahuatldn): Quiexobra, 2300 m, deep rocky gorge, 14 Oct 1995, Hinton, et al. 26122 (TEX); Xianaguilla, 2325 m, 22 Oct 1995, Hinton, et al. 26327 (TEX); above Xianaguilla, 2550 m, 23 Oct 1995, Hinton, et al. 26357 (TEX); San Francisco Ozolotepec, 2815 m, 8 Aug 1996, Hinton, et al. 26839 (TEX); 35 mi. SE of Ejutla, road to Puerto Angel, 22 Aug 1980, Turner 80A-9 (TEX). All of the above collections are said to be common shrubs or trees, varying from 2.0-5.0 m high and collected between 2240-2815 m in pine-oak forests. The closest relative of var. miahuatlana is the allopatric var. subvillosa, which is consistently described as a suffruticose herb or shrub 0.5-2.0 m high, occurring in pine-oak forests from 2100-2500 m. In Crawford’s (1970) treatment of the Coreopsis mutica complex, var. miahuatlana, because of its 5-6 ray florets, will key to or near C. mutica var. microcephala D. Crawford, a taxon with smaller heads occurring to the east of the Isthmus of Tehuantepec. I gave this identification to the first collections of the present plants obtained from the Miahuatlén area by James Hinton, but the fine series of subsequent collections obtained by this ardent field botanist has lead me to conclude that the taxon concerned is deserving of formal recognition. ACKNOWLEDGMENTS _I am grateful to Gayle Turmer for the Latin diagnosis, and to her and Justin ae for reviewing the manuscript. Ms. Maria Thompson provided the illustration. 294 Pit ¥ TOLGGIA Apnmil 1996 volume 80(4):291-294 LITERATURE CITED Crawford, D. 1970. Systematic studies on Mexican Coreopsis (Compositae), Coreopsis mutica: flavonoid chemistry, chromosome numbers, morphology, and hybridization. Brittonia 22:93-111. . 1981. A new variety of Coreopsis mutica (Compositae) from Mexico. Bnittonia 33:547-554. Tumer, B.L. 1992. Coreopsis mutica var. guerreroana (Asteraceae), a new taxon from México. Phytologia 73:7-13. Phytologia (April 1996) 80(4):295. NOMENCLATURAL CORRECTION IN VIOLA (VIOLACEAE) R. John Little University Herbarium, University of California, Berkeley, California 94720 U.S.A. ABSTRACT Viola sororia Willd. var. affinis (Leconte) L.E. McKinney was incorrectly used as the basionym in a previously published combination to change this taxon to subspecific rank. This note corrects the previous combination. KEY WORDS: Viola, Violaceae, nomenclature Viola sororia Willd. var. affinis (Leconte) L.E. McKinney (McKinney 1992:39), was used incorrectly as the basionym to change this taxon to subspecific rank (Little 1992). The following nomenclatural change is made to correct this error and to achieve consistency with the treatments of Violaceae in the Jepson Manual Project and for the Vascular Plants of Arizona Project: Viola sororia Willd. subsp. affinis (Leconte) R.J. Little, stat. et comb. nov. BASIONYM: Viola affinis Leconte, Ann. Lyceum Nat. Hist. N.Y. 2:138. 1826. ACKNOWLEDGMENTS I thank Dr. Bruce Bartholomew of the Botany Department, California Academy of Sciences, Golden Gate Park, San Francisco, CA, for bringing to my attention the need to correct the basionym reference and for reviewing this note. LITERATURE CITED Little, R. J. 1992. Nomenclatural changes in Viola (Violaceae). Phytologia 72:77- 78. McKinney, L.E. 1992. A taxonomic revision of the acaulescent blue violets (Viola) of North America. Sida, Bot. Misc. No. 7. 295 Phytologia (Apnil 1996) 80(4):296-327. NEW LOCATIONS AND INTERPRETATION OF VERNAL POOLS IN SOUTHERN CALIFORNIA Richard E. Riefner, Jr. Department of Parks and Recreation, Orange Coast District, 3030 Avenida Del Presidente, San Clemente, California 92672 U.S.A. & David R. Pryor Department of Parks and Recreation, Orange Coast District, 18331 Enterprise Lane, Huntington Beach, California 92648 U.S.A. ABSTRACT An undocumented series of disturbed and remnant vernal pools was discovered during the abnormally wet 1992-1993 rainfall season in coastal southern California at San Clemente State Beach in southwestern Orange County, and from two sites at San Onofre State Beach in northwestern San Diego County. An additional series of remnant terrace vernal pools was found between Rancho Laguna and the City of San Clemente also in coastal Orange County. San Diego pools associated with either claypan or duripan soils have different suites of plant species despite their close proximity. The coastal Orange County pools, herein proposed as possible liquefaction-ongin pools, also support a distinctive flora. Aerial photographs of the region made prior to extensive urbanization show widespread fields of mima mounds (which coincide with present pool localities), likely indicating extensive historical vernal pool habitat. In coastal Orange County, remnant mima mound-type topography is present only at San Clemente State Beach and at Moulton Meadows in Laguna Beach; fragmented mounds occur in the Dana Hills and the City of San Clemente. We suggest that the origin of some mima mounds in coastal southern California may be attributed to Holocene seismic activity and liquefaction-related events. A biogenic maintenance hypothesis 1s proposed for the mima mounds: foraging activities by fossorial (burrowing) rodents in liquefaction terrain may play an important role in deterring long-term erosion of mounds and filling in of intermound pools. New word models are presented to explain how these relationships might have evolved including: “earthquake successional sequences” and “seismogenic tracker species.” Earthquake successional sequences, including intermittent pools and mounds, provide habitat created by 296 Riefner & Pryor: Southern California vernal pools 297 ongoing seismic and liquefaction-related activities for invasion by seismogenic tracker organisms, which possibly include fairy shrimp, western spadefoot toad, pocket gophers, and other opportunistic species whose mode of life is in part associated with seismic-induced terrain. We describe the previously unrecognized role of cryptogamic vegetation in regulating microscale gradients such as moisture status, nutrient availability, soil temperature, microbial processes, and safe-site seed selection which directly affect the biology of vernal pools. The cyanobacterium, Microcoleus vaginatus, is identified as a potential keystone organism occupying vernal pools and astatic alkaline sinks in southern and central California) A preliminary catalog of the cryptogamic and phanerogamic flora of vernal pools and adjacent upland terrain is presented. A number of rare or regionally uncommon vascular plants were collected during this study including Atriplex pacifica, Dudleya blochmaniae, and Myosurus minimus. New records documented for Orange County include the native Trifolium variegatum and the exotics Afriplex lindleyi and Hainardia cylindrica, Glinus lotoides, introduced from Europe, is new for San Diego County. KEY WORDS: Vemal pools, California, Orange County liquefaction-ongin pools, San Diego County dunpan/claypan pools, fossorial rodents, mima mounds, Holocene earthquakes, seismogenic tracker species, cryptogamic crusts, Microcoleus vaginatus, keystone species, fairy shrimp INTRODUCTION California’s vernal pools are small to medium sized temporary ponds which form above hardpan, claypan, or volcanic mudflow soils during the winter rain season but drain completely by late spring (Bauder 1986); vernal pools are often defined and identified by their endemic or regionally restricted flora (Thorne 1984). They range in size from a few square meters to several hectares and are always shallow, most by 10 to 60 centimeters (Zedler 1987). Pools larger than 20 hectares are referred to as vernal lakes (Holland 1976). Vemal pools are also surface water depression wetlands according to Novitzki (1979). Perched above an impermeable soil layer and separated from groundwater or stream channel inflow, a vernal pool fills only by slowly collecting precipitation, although the wet-life of vernal pools may be extended by subsurface flows (Hanes, ef al. 1990). Although evaporation exceeds precipitation in most coastal southern California wetlands (Stevenson & Emery 1958), vernal pools also share characteristics of ombrotrophic (rainy) environments since precipitation- dominated wetlands receive nutrient inputs primarily from wet and dry atmospheric deposition (Doss 1995). Despite the recognized importance of hydrogeologic regimes in driving nutrient dynamics in wetlands (Carter 1986; Labaugh 1986), the relationship lee hydrologic processes and nutrient cycling in vernal pools has not been addressed. Zedler (1990) has likened the winter vernal pool habitat to a cool-temperate pond, and in late spring to autumn, a desert. Nestled within a subtle upland depression, the pool erratically fills and empties, often repeatedly throughout the season due to uncertain rainfall before completely drying by evapo-transpiration. The dry vernal 298 PHYTOLOGIA Apnil 1996 volume 80(4):296-327 pool pan, also described as a dry marsh bed (Kopecko & Lathrop 1975), may persist for years without filling completely, especially during severe drought. Winter ponding deters most aggressive upland plants from invading a vernal pool; and in shallow pools, the relatively short-lived inundation period also slows development of the permanent anaerobic conditions necessary for establishment of many marshland species. The recurrent inflow of rainwater produces an ephemeral amphibious environment with micro-niches for vascular plant and cryptogamic species including an easily overlooked, shallow undulating pool margin that simulates a tidal mud flat. Alternating winter inundation and summer drought has selected a unique assemblage of amphibious plants and animals including specialized endemics as well as a group of aquatic generalists capable of tolerating a double-stress regime. Zedler (1990) developed the “recurrent gap” hypothesis based upon the reliable recurrence of microhabitats due to uncertain rainfall to explain traits in annual seed plants; accordingly, we emphasize the importance of the “recurrent flux” of inundating rainfall in the creation of niches for cryptogams in vernal pool terrain. The floristic composition of a vernal pool may be perpetuated for decades because southern California’s arid climate does not support peat formation, a process that speeds up plant community succession; the amphibious character of this -habitat and its specialized biota are closely dependent on California’s Mediterranean-type climate which is characterized by mild, moderately wet winters and rain-free summers. Zedler (1987) describes four stages of pool development: 1) a wetting phase; 2) an aquatic phase; 3) a drying phase; and 4) a drought or dormant phase. Vemal pool plants and animals often depend upon one or more of these specific phases (Zedler 1987; Holland & Jain 1984). Although Zedler (1987) indicates that climate is key for the establishment of true vernal pool habitat, edaphic factors may be more important with respect to the vegetation than climate (Holland & Dains 1990). Vermal pools are not unique to California (Thome 1984), but their plant communities of course are (Holland & Jain 1977). Intermittent pools and pool- forming processes that most closely simulate and support biota characteristic of California’s vernal pools, occur primanily in other Mediterranean climate regions (i.e., South Africa, southwestern Australia, the Mediterranean Basin, and Chile). Phytogeographic phenomena, including amphitropical disyunctions of the same or similar phanerogamic species between the pools of temperate North and South America, also link the vernal pool biota of the Mediterranean climate regions. Long- distance dispersal by migrating birds is probably involved (Raven 1963). Members of California’s vernal pool flora which have disjunct austral counterparts include species of Marsilea, Crassula, Isoetes, Myosurus, and Eryngium (Zedler 1987). New information on the phytogeography of cryptogamic vegetation suggests that species shared between southern California and other Mediterranean climates may be relicts of an ancient Madro-Tethyan flora rather than products of long-distance dispersal, which is less likely for many lichens and bryophytes. The cryptogamic flora of South Africa and California, unlike the vascular flora, does share identical and vicariant taxa, i.e., Buellia halonia (Ach.) Tuck., Punctelia punctilla (Hale) Krog, and Trichoramalina crinitum (Tuck.) Rundel & Bowler/T. melanothrix (Laur.) Rundel & Bowler (Bowler & Rundel 1974; Riefner 1989; Weber 1993). The recent discovery of Ramalina canariensis Steiner in California (Riefner & Bowler 1994), previously documented from all the Mediterranean climate regions except California, points to our lack of understanding of dispersal mechanisms and relict endemism on the continental level. Riefner & Pryor: Southern California vernal pools 299 Additional study may yet link the cryptogamic vegetation of California’s vernal pools with that of other Mediterranean climates. Vernal pools and much of their biota are considered to be the most specialized and endangered type of wetland in California (Bauder 1986; Cheatham 1976, Cochrane 1985; Ferren & Pritchett 1988; Ferren & Fiedler 1993; Holland & Griggs 1976; Thorne 1984; Zedler 1987). The rigors of the habitat have provided fertile ground for rapid evolutionary radiation in the vascular flora (Stone 1990). Noteworthy are several characteristic genera of annual plants including Downingia, Eryngium, Lasthenia, Limnanthes, Navarretia, Orcuttia, Plagiobothrys, and Pogogyne. Many taxa often belong to a suite of narrowly restricted species which occupy a particular pool type characterized by specific soil properties, inundation regimes, and/or elevation (Bauder 1993). The likelihood that a particular suite of species is present in any one pool group is determined by a highly localized set of gradients that may not be widely encountered. Bauder (1993) has discussed differences in microscale gradients of soil and elevation affecting vernal pool plants in San Diego County, including highly localized species such as Pogogyne abramsii J.T. Howell and Downingia concolor E. Greene subsp. brevior McVaugh. Similar evidence documenting the microscale conditions of soils and inundation regimes for specific suites of species in Merced and Placer counties was presented by Holland & Jain (1990). Selective pressures exerted by multiple local and microscale parameters including soils, inundation regimes, temperature, and factors such as fungal associates which may affect germination have contributed to the complexity of vernal pool ecology and evolution. Holland & Jain (1990) believe that floras of vernal pools differ according to soil type, while Bauder (1987) and Zedler (1987) state that local pool composition and structure in southern California are best explained by frequency and duration of ponding. We believe a more intricate web of functionally linked physical- hydrogeologic and biological processes including local hydrologic cycles and climate patterns, landform age and origin of mound-depression microrelief, nutrient cycling, microbiota and cryptogamic vegetation, abiotic and biotic factors which maintain mound-depression terrain, site history, and disturbance all interact to affect the diversity and abundance of plants and other organisms in southern California’s vernal pool terrain. Vemal pools occur within an upland vegetation matnx of grassland, chaparral, coastal sage scrub, and/or oak woodland habitat. Because of their isolation, vernal pools have frequently been referred to as terrestrial or ecological islands (Holland & Jain 1988; Zedler 1987; Schoenherr 1992; Stone 1990). The quasi-endemic nature of vernal pool “island plants” is due in part to lack of seed dispersal mechanisms which favors keeping the seed supply on-site. Zedler (1990) compiled data indicating that nearly three-quarters of the vernal pool species in San Diego County have no obvious means of dispersal. Clusters of “island” pools have been referred to as archipelagoes, suggesting that principals of island biogeography might be used to unravel complex phytosociological patterns correlated with pool numbers and size (Holland & Jain 1981, 1988). In this paper, we refer to local patches of remnant pool habitat as islands within an upland vegetation matnx, and to chains of such patches as archipelagoes (Burkey 1995). Herein, we report new localities of San Diego claypan and duripan vernal pool 300 PHY TOLOGIA Apnil 1996 volume 80(4):296-327 habitat and a new archipelago comprised of the previously uncharactenzed Orange County vernal pools. The San Diego and Orange county pools each support different suites of plant species. The ready accessibility of these pools with their distinctive floras offers a rare opportunity to study the complex, and stil incompletely known, biotic interactions within vernal pool communities. DISTRIBUTION OF CALIFORNIA VERNAL POOLS In western North America, vernal pools have been reported from southern Oregon southward into Baja California, México (Holland & Griggs 1976; Kagan 1986; Moran 1984). The vernal pools of California primarily occur in two clusters: 1) the coastal terraces and areas of gentle topography of the lower coastal mountains from Sonoma to San Diego County; and 2) from Shasta to Kern county in the Central Valley (Zedler 1987; Holland & Jain 1977; Ferren & Pritchett 1988). Vernal pool vegetation is best developed in the eastern Central Valley on ancient terrace soils bordering the foothills of the Sierra Nevada, where concentric rings of wildflower displays: unmistakably reveal the presence of vernal pool terrain (Holland & Jain 1988). Holland (1978a; 1978b) states that vernal pools were apparently a common feature in most of the Central Valley in presettlkement times, estimating that nearly one-third of the valley historically supported them. However, vernal pools in other regions of the state have not been studied as thoroughly, so we cannot adequately assess their historic range and estimate habitat loss. In southern California, Ferren & Fiedler (1993) estimate that as much as 90% of the vernal pools have been destroyed in the past century. In San Diego County, vernal pools were once common on coastal terraces and inland valleys (Purer 1939). Many have been destroyed by development. Current estimates indicate over 93% of the county’s vernal pool habitat has been extirpated, and many of the remaining pools are highly disturbed (Bauder 1986). The situation is similar in Orange County. Aenal photographs of northwestern San Diego and southwestern Orange counties taken prior to extensive urbanization (1932 Whittier Collection; 1941 USGS; 1953 USDA; 1964 California Coastal Commission) show widespread fields of mima mounds, indicating extensive historical vernal pool habitat. Vernal pools are well known in San Diego and Riverside counties, but not in Orange County, where the pools have remained undocumented. This overlooked habitat has been omitted in important studies of cniical plant communities, such as Horn, et al. (1993) and Sawyer & Keeler-Wolf (1995). Evans & Bohn (1987) identified mima mound topography in southern Orange County, and Marsh (1992) described remnant or extirpated pools from the Laguna Beach area and Dana Point. Marsh (1992), however, did not cite any of the vernal pool obligate species discussed by Zedler (1987) and therefore, could not authenticate the presence of true vernal pool habitat. Marsh (1992) believes that vernal pools were once common in southern Orange County. Remnant Orange County terrace vernal pools were identified during this study utilizing aerial photography provided by G. Kuhn (1993, pers. collection), soil survey, and herbarium specimen data. Several pools were documented between Riefner & Pryor. Southern California vernal pools 301 Rancho Laguna and the City of San Clemente only during the unusually wet 1992- 1993 rainfall season (Orange County Environmental Management Agency 1996). Aerial photographs taken prior to urbanization in south-central Orange County (City of Irvine, University of California Campus) depict vernal pools (1971, W. Bretz-NRS Collection). Other vernal pools recently recorded from Orange County occur at Whiting Ranch and O’Neil Park (Jones & Stokes 1993), and in Fairview Park, Costa Mesa, (Bowler, et al. 1995). A series of small pools also occur near the Badlands Park in Laguna Beach (P. Bowler, pers. comm.). The remaining vestige of pools in Orange County are possibly remnants of a once broader complex of coastal terrace habitats that has been nearly extirpated because of urbanization and agriculture. Soil surveys show that vernal pools in coastal southern Orange County are developed primarily over the very slowly permeable, moderately alkaline Myford sandy loam or the moderately slowly permeable Botella clay loam (Soil Conservation Service 1978). A map and detailed study of the flora of these pools is in preparation. THE ORIGIN OF MIMA MOUNDS Mima mounds are the elevated, often circular areas between vernal pools that are composed of unconsolidated fine soils; the term mima mounds originates from their type locality, Mima Prairie, near Olympia, Washington (Dalquest & Scheffer 1942). Mima mounds and vernal pools are inexorably intertwined; there are no mounds without intervening depressions. In California, the intermound depressions are also known as hogwallows (Arkley & Brown 1954; Brandegee 1890; Cox 1984a). Although climate is key for the formation of vernal pools, topography is also important, since pools mostly form in closed depressions (Zedler 1987). In North America, mima mounds are recorded only west of the Mississippi River ranging from southern Canada to northern México (Cox & Scheffer 1991). Studies of mima mound formation have produced a number of controversial theories about the origins of the mounds. Washburn (1988) provided a comprehensive review, and Berg (1990) grouped theories into four categories: 1) depositional; 2) erosional; 3) periglacial; and 4) biological. Cox (1984b), Zedler (1987), and Holland & Jain (1988) described mound-building processes pertinent to California including origin by: 1) wind deposition near the base of shrubs; 2) groundwater pressure; 3) the activities of fossorial rodents; 4) fracture patterns in the underlying hardpan; 5) expansion and contraction of clay minerals upon wetting and drying; and 6) subsurface mass movements such as soil-piping. Zedler (1987) proposed that differential weathering and settling is primarily responsible for mounded topography in California and presented a model for mound origin by weathering. Each one of the foregoing theories can be locally confirmed by data from particular localities, but none hold true for all situations (Holland & Jain 1988). In recent years, however, only the fossorial rodent hypothesis has received broad acceptance (Dalquest & Scheffer 1942; Arkley & Brown 1954; Barry 1981; Cox 1990). According to the fossorial rodent hypothesis (Cox 1984; Cox & Scheffer 1991), moundfields originate in shallow soils where pocket gophers build nest sites to escape predation and weather. The nest sites become the center of fixed territories. Subsequently, gophers mine and translocate soil in slow centripetal fashion toward 302 PHY:FOLGELIA Apnmil 1996 volume 80(4):296-327 nests. This tunneling translocates soil, which in time, builds mounds until the centripetal soil-mining activities are balanced by unknown factors (Cox & Scheffer 1991). Cox (1984a,b) has provided evidence that mound formation by foraging rodents is plausible; estimating a typical mound could be formed in approximately 108 years. Extensive behavioral documentation of gophers by numerous researchers has apparently corroborated the pocket gopher theory (Gregory, et al. 1987; Hansen 1962; Howard & Childs 1959). Scheffer (1958) and Cox & Scheffer (1991) believe that mima mound terrain occurs only where rodents are, or have been, working the soil. Although there is an extensive literature discussing the biogenic origin of mounded landscapes, geologists have remained skeptical. Berg (1989) proposed that mounded landscapes are formed when strong seismic activity occurs in areas where a shallow mantle of loess or other fine unconsolidated material overlies a relatively ngid, planar substratum. The substratum could be hardpan, bedded gravel, or bedrock. Berg (1990) then presented data showing that the circular shape and the uniform pattern of mounds can be explained by Richter’s (1958) theory of seismic wave motion. According to Berg (1990), the distribution of mima mounds in the United States is directly correlated with regions of moderate to high seismicity. The seismic hypothesis could account for the presence of mima mound-type topography in a wide variety of global geomorphic and climatic provinces and could also explain mound uniformity and soil profiles of mima mound landscapes that cannot be easily demonstrated by the pocket gopher or other theories. Recently, Kuhn, et al. (1995a) identified liquefaction-related features including fissure fills and lateral spreads in mima mound terrain near Carlsbad, San Diego County, in coastal southern California. Paleo-liquefaction, as postulated by these authors, is conspicuous at the Carlsbad site as widespread and abundant injection dikes composed of fine grained white sand, inferred to be ancient beach sand, thrust through the plastic clay-rich surficial deposits; the injection dikes are “sand blow deposits,” previously described by Fuller (1912) and Sieh (1978). Existing sand blow deposits (mima mounds) are correlated with the extent of an ancient intertidal lagoon. Kuhn, ef al. (1995b) hypothesize that during or since the Holocene, mima mound formation due to liquefaction only occurred rarely, when a strong earthquake coincided with unusually high rainfall and perched water conditions far above modem groundwater levels. Although the age of the liquefaction event(s) remains to be determined precisely, the size of the sand blow deposits, the area affected, and the uplift of ancient intertidal deposits imply that coastal uplift and mima mound formation occurred as a result of large, infrequent pre-historic earthquake(s) of magnitude seven or greater. Legg, et al. (1994) and Kuhn, et al. (1995a) suggest that the source of seismic activity may well be the Newport-Inglewood/Rose Canyon fault located four to six kilometers off the southern California coast. Later studies by Kuhn, et al. (1995b) have identified paleo-liquefaction features which extend upward into a series of regressive continental deposits that overlie flights of marine terrace platforms ranging in elevation from 10 to 60 meters. Cross-cutting stratigraphy and relative weathering show at least three epochs of paleo-liquefaction in this region that have displaced very old Indian middens and other archaeological sites (Kuhn, et al. 1995b). Thus, major Quaternary deformation of the southern California coast induced during large strike-slip earthquakes are recorded by liquefaction features which could be important in the formation of vernal pools located on higher terraces in Orange and San Diego counties. Riefner & Pryor: Southern California vernal pools 303 Norwick (1991) described the relationship between vernal pool formation and geomorphic processes. He identified ongoing tectonic activity, including development of surface folds and shutter ridges (when a fault moves rock masses horizontally across a valley) with the formation of sag ponds, vernal pools, and swale topography. Norwick’s “sand volcanos” and liquefaction craters described from the San Andreas Fault zone were formed during the earthquake of 1906, and apparently predate other intermittent pools of the region. Sieh (1978) also identified liquefaction-related features, including sand blow deposits, as by-products of slip along the San Andreas fault. Previously, Fuller (1912) studied the succession of shocks collectively designated as the New Madnid earthquake, which occurred between 1811 and 1812, in the central Mississippi Valley in an area encompassing portions of Missouri, Arkansas, Kentucky, and Tennessee. No other feature of the New Madrid region is so conspicuous or widely encountered as the “sunk lands” which resulted from local settling or warping of alluvial deposits (Fuller 1912). Sunk lands are characterized by major alterations to or creation of wetland habitats in clayey alluvial deposits that were still evident 100 years following the shocks. These include sand sloughs, river swamps, sinks, lakes, and ponds. Fuller also describes in great detail sand blows and fissures, which are presently interpreted as relict features of liquefaction events (Obermier 1989). Fuller cites numerous historical accounts of fields composed of sandy mounds, including (pg. 81): 1) “In several places the [sand] blows so obstruct the drainage as to cause the water to collect in shallow pools throughout the wet season.”; 2) “The [sand] blows were so thick as to touch, giving rise to many irregular depressions, in some of which considerable pools of water accumulate.”; and (pg. 83) 3) “The country here was formerly perfectly level and covered with prairies of various sizes dispersed through the woods. Now it is covered with slashes (ponds) and sand hills or montecules, which are found principally where the earth was formerly the lowest... .” The sand blow regions coincide with Berg’s (1990) regions of past moderate to high seismicity. Fuller also documents sand craterlets formed near San Francisco in the earthquake of 1906. The seismogenic origin of mounded landscapes proposed by Berg (1990) and Kuhn, et al. (1995b) works also for coastal southern California. It is plausible that the origin of mima mound terrain is, in part, a product of paleo-liquefaction induced by large pre-historic earthquakes, possibly occurring since the Holocene. The impact of shaking on mound-pool terrain could produce settling of fine mound sediments even when earthquakes are not powerful or when they do not coincide with perched water conditions. If mima mounds in coastal southern California were originally formed by successive episodes of liquefaction (presumably southern California has been subjected to large seismic events which induced mound-pool formation long before the Holocene), what ongoing factors operate to maintain the circular profile and height of mounds and deter siltation of fine sediments and sand into intermittent pools? We Suggest a biogenic maintenance of liquefaction-mounded landscapes by the activities of fossorial rodents. As discussed above, evidence has been accumulating for decades indicating that pocket gopher activity is associated with mima mound terrain (Dalquest & Scheffer 1942; Arkley & Brown 1954; Barry 1981; Cox 1990). Soil mining and translocation within “mima mounds” by pocket gophers could replace fine sediments easily lost to erosion, accounting for the long-term maintenance of the mounds’ circular form and 304 PHY TOLOGIA Apnil 1996 volume 80(4):296-327 height. Tunneling and soil translocation may also impede the otherwise inevitable siltation of intermound basins. Cox (1984b) believes the activities of fossorial rodents create intermound basins while Kuhn, et al. (1995a) state that some vernal pools originate over liquefaction-related laterally spreading clays. Kuhn (unpublished data) has observed vacant rodent burrows in liquefaction terrain in southern California. Abandoned or declining gopher populations in mima mound terrain could produce a mosaic of swales and cloudy silt-pools that provide favorable habitat for other organisms including fairy shrimp, but not for vernal pool obligate plants. Some species of fairy shrimp are not randomly distributed in natural temporary ponds but may favor pools with turbid water to avoid Notonectid (i.e., backswimmer) predation (Woodward & Kiesecker 1994). Shallow silt-pools lack inundation regimes necessary to sustain year to year vernal pool obligate plants except during abnormally high rainfall years. However, cloudy later succession “seres” of intermittent pools could be important in the distribution of vernal pool plants and other species which lack long-distance dispersal mechanisms by providing local habitat patches in the chain of biological archipelagoes. Seed and other propagules could be dispersed between vernal pool “islands” by birds (Baker, et al. 1992). Study of liquefaction terrain could afford insight into abiotic and biotic processes affecting possibly coevolutionary biological interactions in earthquake landscapes. The widely accepted hypothesis that the ongin of mounded topography and associated intermittent pools in California is the result of only a single or simplified process should be reevaluated. Vitek (1978) also concluded that mounded topography may result from various processes, and Fuller (1912) described the differences in the genesis of sand blow mounds and prairie mounds. In California with its complex geologic history, mound-depression landscapes can be attributed to numerous and often complementary processes, so that interacting biotic and abiotic processes need to be quantified at each site. In coastal southern California on high geomorphic surfaces (pre-Holocene), mound topography could have a complex ongin; mounds may have formed when the surfaces were at low elevations, particularly in sandy sediments conducive to liquefaction, but after uplift, the mounds would have been modified by fossorial rodent activity (R. Shlemon, pers. comm.). Mima mounds and pools developed on specific geologic formations or soil series with different inundation regimes, water-retention capacities, stability and sediment shedding characteristics would be expected to support distinctive floras and faunas. Liquefaction-origin pools developed in sandy substrates are prone to wetted-clay slip and external drainage; human disturbance could easily reduce the frequency and duration of ponding in vernal pools. As a result, liquefaction-terrain vernal pools may not be recognized readily and their accompanying biota could be overlooked; clearly, this has been the case in Orange County. Mima mounds composed of white or tan fine sands occurring in proximity to major fault zones or in soils without an argnilic horizon, especially those on coastal terraces or inland dunes, may be suspected of liquefaction origin (G. Kuhn, pers. comm). Aerial photographs depicting white, circular spots may indicate remnant mima mounds (i.e., sand blow features) that have been mechanically bladed or disced (Kuhn, et al. 1995a). Vemal pools on uplifted marine terraces in the San Diego area previously discussed by Purer (1939) should also be inspected for liquefaction features. Riefner & Pryor: Southern California vernal pools 305 FLORISTICS AND ECOLOGY OF CRYPTOGAMIC VEGETATION IN VERNAL POOL TERRAIN One of the most widely discussed topics of California natural history is the distinct, concentric assortment of vascular plant species which describe subtle topographic gradients in vernal pools. In southem California the floras of vernal pools differ (Bauder 1987), but whatever species are present selectively inhabit micro-niches or broad radial zones related to inundation regimes, and perhaps, nutrient availability. Kopecko & Lathrop (1975) describe five habitat zones for vascular plants in vernal pools on the Santa Rosa Plateau in western Riverside County, but cryptogamic species also display this kind of zonation. Cryptogams in vernal pools have scarcely been studied in California, and their contribution to the ecology of this ecosystem has been severely neglected and warrants immediate attention. A preliminary survey of southern California vernal pools suggests a cryptogamic community dominated by cyanobacteria, bryophytes, and cyanophilous lichens (lichens that have blue-green algae photobionts) occupy zones similar to those of the specialized vascular plants. Kopecko & Lathrop’s (1975) “muddy margin zone” and the “vernally moist zone” are occupied and often temporarily dominated (extending from the pool basin to mesic margin) by such organisms including species of Nostoc (cyanobactennum), Scleropodium (moss), Microcoleus (cyanobacterium), Fossombria (liverwort), Collema (cyanolichen), Riccia (liverwort), Funaria, Bryum, and Ceratodon (mosses). Nostoc, Leptodictyum (moss), Microcoleus, (?) Fissidens (moss), and Scleropodium often thrive in the “vernally standing water zone” of vernal pools, and species of Bryum, Funaria, and Microcoleus are common invaders of the “drying marsh bed.” A narrow, ephemeral band of bryophytes dominated by liverworts which occurs between the “muddy margin and vernally moist zones” indicates seasonal or year to year fluctuations in precipitation. Lichens most frequently become a prominent feature in the “dry grassland/scrub zone” of undisturbed vernal pool landscapes and may include species of Acarospora, Catapyrenium, Collema, Cladonia, Psora, Toninia, and Trapeliopsis. The lichens are successionally dominant to mosses and even certain grass and forb species in specialized habitats (Coker 1966; During & Van Tooren 1990; Watt 1937), such as southern California coastal cliffs, dunes, volcanic tablelands, and littoral communities, but are not dominant in inundated areas of vernal pool landscapes. Later successional bryophyte species and cyanophytes such as Microcoleus vaginatus (Vauch.) Gom. are also important in native grasslands and scrub in southern California. The absence of cryptogamic flora from suitable habitats in and around vernal pools indicates disturbance and/or alteration of hydrologic cycles by human activities including habitat fragmentation, grazing and discing, alteration of the fire ecology, and air pollution (Bowler & Riefner 1990). It should be noted that pristine floras such as the Morro Bay region in San Luis Obispo County support nearly 375 lichen taxa (Riefner, in prep.); the depauperate list of cryptogams presented in this paper is typical of urban environs in coastal southern California. Cryptogamic soil crusts (also called biological, cryptobiotic, organogenic, and microphytic crusts) are formed by complex communities of several groups of microphytes including mosses, lichens, liverworts, fungi (including mycorrhizal fungi), green algae, cyanobacteria (blue-green algae), and bacteria. Soil crusts are 306 PHYTOLOGIA Apnil 1996 volume 80(4):296-327 common in arid and semi-and landscapes, and when well-developed, cover the ground with an almost continuous sheet of photosynthetic machinery equivalent to a layer of phanerogamous leaves (Lange, et al. 1994). Although usually not thicker than a few millimeters, soil crusts play a decisive role in the functional ecology of arid ecosystems, including cycling of nutrients (especially nitrogen), soil aggregation and stabilization, carbon gain in large areas (Eldndge & Greene 1994; Lange, ef al. 1992; Beymer & Klopatek 1991), and soil fertility and microbial food webs (Harper & Marble 1988; Johansen 1993; Belnap & Gardner 1993; Belnap 1995). More importantly, cryptogamic crusts deter surface evaporation by sealing the soil surface, and improve overall moisture storage (Lange, et al. 1992). During rains, these crusts produce a high yield of runoff percolation which does not occur if crusts are broken or absent (Yair 1990; Lange, et al. 1992). Claims of reduced erosion and improved water relations due to cryptogamic vegetation have been widely reported in the literature, but until recently, the evidence has been largely circumstantial. Several authors, however, have recently described and expenmentally reproduced the mechanisms by which cryptogams protect the soil and regulate water flow (Chartes & Mucher 1989; Eldridge & Greene 1994; Tchoupopnou 1989; Kinnell, et al. 1990; Yair 1990). Thus, the presence or absence of cryptogamic crusts in arid regions can influence the hydrologic cycles of total landscapes including soil erosion (Cameron & Blank 1966; St. Clair & Johansen 1993; West 1990). Despite their importance and widespread distribution, our present knowledge of the species composition and mode of life of cryptogamic crusts is extremely limited in California where research has largely focused on the study of higher plants. In undisturbed vernal pool landscapes cryptogamic crusts are often a prominent feature that are easily destroyed by grading, discing, grazing, alteration of the fire ecology, and trampling. In a dry climate, human alteration of gently mounded topography characteristic of vernal pool terrain can easily disrupt the delicate balance of specialized hydrophytes inhabiting the narrow radial zones of shallow pools. Additional loss of rain inception and runoff generated percolation historically produced by cryptogamic crusts may dramatically influence the hydrologic status of disturbed vernal pools in southern California, where the average rainfall is markedly less than in the northern part of the state. Increased erosion and decreased aggregate soil stability associated with degradation of cryptogamic communities of disturbed sites (Eldridge & Greene 1994; Kinnell, et al. 1990) could also negatively affect the biology of vernal pool terrain. The impact of severe erosion in vernal pool terrain is likely to be greater due to increased transport of organic nitrogen by eroded sediments where the erosional products are nutrient-rich silt and clays rather than sand (Burwell, et al. 1975; Pallis, et al. 1990; Kinnell, et al. 1990). Disturbance of soil surface crusts can also affect vascular plants due to disruption of food webs and alteration of the soil microbiota including mycorrhizal and rhizosheath associations (Allen 1991; Fitter 1977; Grime, ef al. 1987; Hardie & Leyton 1981; Hartnett, et al. 1993), decreased water availability and nutrient uptake in vascular plants (Belnap 1995; Harper & Pendleton 1993; Rogers & Burns 1994), and decreased germination and seedling establishment (Lange, et al. 1994; Harper & Pendleton 1993; Rogers & Burns 1994; West 1990). The absence and/or reduction of cryptogamic crusts throughout much of southern California’s remaining vernal pool landscapes raises many questions critical to the long-term management of this threatened, arid-land resource. The microphyte-rich communities inhabiting southern California's vernal pool landscapes support several interesting, strongly interacting associations occupying the Riefner & Pryor: Southern California vernal pools 307 muddy margin and vernally moist zones which could potentially affect the biology of other pool species. During an ongoing investigation of vernal pools in southern California, observations by the senior author of several undisturbed and disturbed pool complexes indicate the dominant constituent of these crusts, the cyanobacterium (blue- green alga) Microcoleus vaginatus, could influence or directly regulate several microscale gradients which affect seed plants competing for limited resources in narrow zonal boundaries. This paper presents several mechanisms by which cryptogamic crusts dominated by M. vaginatus regulate microscale gradients including water relations, nutrient availability, microbial processes and soil microbiota, soil temperature, safe-site seed selection, and seedling enhancement in vernal pools. Microcoleus vaginatus is a filamentous, black, mat-forming species which secretes a mucilaginous polysaccharide material from a network of filaments which bind other microphytes and fine sediments into a well-developed crust (Belnap 1993a). Microcoleus vaginatus commonly comprises the major component of cryptobiotic crusts in many western states (Belnap 1993a). The success of this cyanophyte in occupying a wide habitat range is due to its resistance to temperature extremes, hypersalinity and alkalinity, desiccation, and to modest nutrient requirements (Carr & Whitton 1982). Microcoleus is ubiquitous, and the crust is often inconspicuous as a dark film in the “dry grassland zone” of Kopecko & Lathrop (1975). Microcoleus, however, develops a conspicuous mat or crust in vernally moist to inundated clay depressions in grasslands, alkaline sinks, and vernal pools if alternating cycles of wetting and exposure are present which favor rapid filament and sheath production. During the aquatic phase of vernal pools, M. vaginatus is often the locally dominant organism forming extensive mats over a broad zone extending from the outer pool edge to standing pool water. As ponded waters fluctuate, the Microcoleus colony simultaneously swells into a motile mat covering exposed mud to reach favored photic zones. During the drying phase of vernal pools as the perimeter retreats, Microcoleus does also, often coinciding with the germination of vascular plants. The desiccating mat forms a crust/shell over saturated pool soil, deterring evaporation and enhancing the life of an anaerobic zone which inhibits upland vascular plants from colonizing pool margins, and importantly, promotes growth of vernal pool obligates by providing fuel in the form of ammonia. After completely drying, “the dry marsh bed” of shallow pools is often richly covered by Microcoleus, and in late spring and summer the curling crusts are a conspicuous feature of many alkaline sinks and vernal pools. This feature could be useful for identifying seasonal wetlands and perched water conditions in southern and central California. Two key ecophysiological characteristics of Microcoleus vaginatus could account for possible mechanisms that effect important microscale gradients. (1) Microcoleus vaginatus produces a large, distinct, sticky extracellular sheath that surrounds groups of living filaments (Belnap 1992). Sheath matenal rapidly absorbs water, and when wetted, swells, then mechanically extrudes through or over the soil; as the substrate dries the filaments secrete additional mucilaginous matenal. Rewetting repeats this cycle. Microcoleus vaginatus frequently develops a mat or drying crust several millimeters thick due to the recurrent flux of vernal pool waters. Microcoleus is capable of growing up to five centimeters in 24 hours (Belnap, ef al. 1993b) when wetted which enables it to reach favorable photic zones along fluctuating or subsiding pool waters. These adaptative mechanisms may maintain the mat in an 308 PHYTOLOGIA Apnil 1996 volume 80(4):296-327 active state in exposed circumstances during alternating cycles of inundation and exposure. (2) A second trait of Microcoleus vaginatus and other cyanophytes is the ability to fix atmospheric nitrogen throughout most of the year (Fuller, et al. 1960). Cyanophytes produce nitrogenase, an enzyme complex responsible for catalyzing the conversion of dinitrogen to ammonia. Nitrogen fixation is a stringent anaerobic process, and since M. vaginatus lacks heterocysts (thick-walled cells that exclude oxygen), an anaerobic microenvironment must be supplemented in other ways for nitrogenase activity to produce ammonia. Belnap, ef al. (1993b) has postulated that M. vaginatus creates an oxygen-free zone by packing multiple filaments within thick extracellular sheaths or packing groups of sheaths together which has been demonstrated by the morphologically similar oceanic species Méicrocoleus chthonoplastes (Paerl 1985, 1990; Paerl, et al. 1991; Pearson, ef al. 1981). Nitrogenase and N-fixation activity may be closely related to growth of new sheath material and/or new filaments produced only when this cyanophyte is wetted (Belnap, unpublished data). Asexual reproduction by mat expansion acts as a nitrogen source and sink which is mobilized by the presence of water. j Several other nitrogen-fixing lichen and cyanobacteria species are present in vernal pool habitats including Nostoc sp. (which does develop heterocysts not requiring constant anaerobic conditions) and the cyanolichen Collema tenax (which hosts Nostoc as its phycobiont). The cyanolichens apparently occur in numbers too small to contribute significant fixed nitrogen into this system, but species of Nostoc may, particularly in deeper static pools. Nostoc, previously discussed by Zedler (1987), is a colonial cyanophyte visible in vernal pools or more frequently along the edge of drying pools as small translucent balls. Nostoc is well documented for salinity tolerance, importance in nitrogen fixation, and use as a biofertilizer (Singh 1961; Singh, et al. 1996). Microcoleus may serve an important function in the population structure of many vascular plants competing in zonal boundaries and microhabitats of vernal pools. The rapid growth and mat/crust-formation of M. vaginatus not only deters evaporation but maintains and enhances an anaerobic environment for itself and associated seed plants in microzones. These affects, in concert with specialized germination requirements, perpetuate native annuals and exclude exotics, and explain the resistance of vernal pools to upland plant invasion. Most importantly, ammonia enrichment produced by Microcoleus could alter competitive interactions and provide fuel for the explosive growth and reproduction of vernal pool annuals. | Numerous studies have demonstrated that the nitrogen fixed by cyanobacteria is available to and is used by neighboring vascular plants (Fuller, et al. 1960; Maryland, et al. 1966; Maryland & McIntosh 1966; Stewart 1967). Also, nutrient uptake of seed plants associated with cyanobacterial crusts has been demonstrated to show higher concentrations of many essential macronutrients (Harper & Pendleton 1993; Belnap 1995). In some ecosystems, these crusts have been demonstrated to be the dominant source of limiting elements for seed plant communities (Evans & Ehlringer 1993). In other studies, Lange (1974) demonstrated that compounds in the gelatinous cyanobacterial sheath material were able to chelate elements essential for growth. Since both clay particles and organic material are negatively charged, the sheath material electrostatically absorbs positively charged essential nutrient ions and holds them in a form available to higher plants (Lange 1976). The sheath material of Microcoleus may also enhance the Riefner & Pryor: Southern California vernal pools 309 availability of iron to vascular plants (Belnap & Harper 1994). Chelation of iron may be especially important in alkaline soils (such as the vernal pool soils within the study region) since iron is usually bound in forms unavailable to seed plants (Wallace 1956). Belnap (1992) also showed that mucilaginous sheath material is often coated with negatively charged clay particles, providing a mechanism for retaining positively charged macronutrients in the upper soil profiles that are otherwise prone to leaching. ‘Swelling Microcoleus mats following the retreating pool-edge waters during the “drying phase” slow evaporation, extend the wet-life of a pool, and may also produce radially-zoned rings of nutrients, inducing certain assemblages of seed plants adapted to specific requirements. A reduction in fecundity in Limnanthes and Orcuttia due to moisture stress (Brown 1976; Griggs 1976) may be moderated by Microcoleus crusts. Linhart (1972; 1974) provided evidence that genetic differences within populations of Veronica peregrina L. are adapted to specific environmental conditions differing between pool edge and basin, and may be attributed to nutrient and competition gradients. Linhart (1976) also documented that Lasthenia and Downingia have greater numbers of viable seed per head at the periphery than at the center of vernal pools. The mucilaginous mat filaments may also act as a seed trap incorporating minute seeds between the polysaccharide sheath material. Lin (1970) noted that certain Limnanthes species are restricted to well-defined, smaller vernal pools with conspicuous radial zonation. Seed size, micromorphology, and omamentation in semi-aquatic plants illustrate the role of safe-site selection (possibly zonation and micro-site establishment) and colonizing ability dependent upon nutrient enriched zonal areas. Thus seed trapping in crusts, and the resulting nitrogen sink, represents an important sequestering of resources which could otherwise be lost in upland areas. The black mats/crusts of Microcoleus vaginatus may also stimulate vascular plant growth and nutrient uptake by producing warmer soil temperatures during the growing season especially at higher elevations or in cooler coastal fog zones. Dark-crusted surfaces have been demonstrated to be significantly warmer than light-colored, non- crusted ones (Belnap 1995). Surface temperatures may be very important, since nitrogenase activity is an extremely temperature dependent process (Rychert, ef al. 1978). Altered soil temperatures can also effect microbial activity and affect plant germination rates and seedling growth, since timing in vernal pool plants is essential for establishment; relatively small delays in germination can reduce species fitness and seedling establishment (Bush & Van Auken 1991). Because pool temperatures tend to follow changes in air temperature differences more closely than other aquatic environments, Microcoleus mats could moderate mud surface temperature fluctuations between pool center and periphery during dry-down (Alexander 1976, Linhart 1976). Consequently, disturbance of Microcoleus mats/crusts can affect moisture status, nutrient availability, seed trapping and germination, seedling establishment, and competitive community structure allied to microzones developed during the recurrent flux of perimeter pool waters. This, in turn, could profoundly affect small annual seed plants in highly competitive vernal pool zones. Since vernal pool annuals produce seeds that germinate in standing water or saturated soils, and most exotic taxa do not, moisture maintenance by specialized crusts is an important resource in excluding exotic plant invasion at the pool ecotone between inundated and non-inundated habitats. Alteration of pool hydrology via reduced precipitation infiltration due to loss of cryptogamic crusts inhabiting open ground characteristic of native grasslands, may also negatively affect subsurface flows and dry-down timing in vernal pools by 310 PHYTOLOGIA Apnil 1996 volume 80(4):296-327 decreasing water storage capacities in certain landscape types, especially in shallow soils. Subsurface flows generated by the surrounding watershed which recharge pools by lateral movement of water are important in southern California (Hanes, et al. 1990; Zedler 1987). Decreased infiltration rates and subsurface flows could negatively alter pool hydrology following the initial pool filling by rainfall. Pools located in native grasslands, i.e., the Santa Rosa Plateau, which support diverse cryptogamic vegetation characteristic of the habitat (Riefner, unpublished data), may have increased water infiltration compared with bare or disced ground. This could significantly influence the recurrent flux of pool waters over the course of the season and contribute to pool diversity/productivity in the more and southern portions of the state. Soil water retention capacity in combination with other biotic, geologic, edaphic, and climatic factors may significantly influence the amount and timing of subsurface flows which are important in supplementing direct precipitation and offsetting evaporative loss from pools (Hanes, et al. 1990). According to terminology presented by Westman (1987), Microcoleus vaginatus would be identified as a “keystone species” (i.e., the addition or the removal of this species could result in marked changes in community structure and function). Paine (1980) proposed that “modules” may exist within a community. Species dependent upon a common suite of resources, which disappear with the removal of a strongly interacting species, or appear with its addition, belong to a module (Paine 1980). Although there have been few detailed studies describing cryptogams as keystone species for vascular plants, a classic example is the moss Sphagnum, which controls the vegetation of bogs at every stage of development by impeding drainage and creating an acid habitat (Crum 1976). Recent research, however, has identified the reindeer lichen, Cladonia rangiformis Hoffm., as an important regulator and a keystone organism of lowland heath communities in the United Kingdom (Newsham, et al. 1995). Quantitative and empirical evidence discussed by Riefner & Bowler (1995) and Knops, ef al. (1991) indicate that the fruticose (bush-like or pendulous) lichens Niebla cerucoides Rundel & Bowler and Ramalina menziesii Taylor improve moisture status and nutrient availability for vascular plant species occupying coastal cliffs and oak savannas (respectively) which concur with Westman’s (1983) speculations that relatively minor differences in moisture availability between habitat sites may be sufficient to select for specific species. Vascular plant species inhabiting vernal pool zones affected by Méicrocoleus possibly include Limnanthes and Plagiobothrys in outer pool margins, and Myosurus and Psilocarphus brevissimus Nutt. which often germinate in drying cyanobacterial crusts following retreating waters. Microcoleus mats/crusts and associated seed plants occupying a broad radial- zone between the edge of the vernally moist zone and the muddy margin of retreating pool waters could be considered a module. The Microcoleus module may affect disturbed pools or clay borrow pits in the way the native nitrogen-fixing lupine (Lupinus arboreus Sims) alters succession by nutrient enrichment of its habitat (Maron & Connors 1996). DISCUSSION We propose that much of the mima mound-type topography in coastal southern California is a geologically young, dynamic landscape which formed as a result of Riefner & Pryor. Southern California vernal pools 311 paleo-earthquake and liquefaction events. Conservation and management of pocket gopher colonies may play a key role in perpetuating vernal pool basins and mima mound terrain in this region. The crater pools described by Norwick (1991), as well as other pools known to have filled rapidly with silt, should be examined for evidence of pocket gopher activity to support the validity and role of the biogenic maintenance hypothesis. Vernal pool obligate plant species may gradually succumb to altered hydrologic regimes due to increased siltation in uncolonized mound-depression landscapes or in terrain abandoned by gophers. Other successional stages of liquefaction-origin pools, however, provide habitat for plant communities associated with later stages of pool formation and siltation and other organisms such as fairy shrimp. Cloudy, successional silt-pools also deter growth of certain cyanobactena, including Microcoleus, which require clear water (J. Belnap, pers. comm.); thereby resulting in decreased nitrogen and water storage capacities which promote further changes in the floristic composition. Intermediate stages of siltation may be evident when vernal pool obligates appear only during abnormally high rainfall years which temporarily restore the inundation regime necessary for germination. Orcutt (1887) described a similar phenomenon in San Diego which occurred only in the unusually wet spring of 1884 when previously dry hollows and flats produced a luxunant array of vernal pool species which “withered away to let others succeed when another favorable season should chance to roll around in future years.” Earthquake successional sequences and seismogenic tracker species, that may include pocket gophers and fairy shrimp, potentially illustrate complex issues of population coevolution which remains a relatively unexplored topic in vernal pool science. The occurrence of certain locally abundant bulb-forming plants, including species of Brodiaea and Muilla, near vernal pools is often attributed to substrate preference, but may be in part due to gophers eating and storing plant parts. Previously, Brandegee (1890) discussed the “rooting of hogs” as a possible means of dispersal of Dodecatheon species locally distributed about vernal pools in Sacramento County. Although the productivity of cyanobacterial mats has been described for other habitats experiencing alternate wetting-drying, i.e., tidal flats (Zedler 1980), the role and potential importance of these mats in vernal pools has not been previously described. Microzone formation by Microcoleus vaginatus which ensures an anaerobic environment for itself and the ability to fix atmospheric nitrogen brings about a dynamic interplay between moisture storage, nitrogenase and ammonia production, and soil temperature patterns which promote rapid growth and reproduction of annual seed plants in vernal pools. Previously, Jokerst (1993) also speculated that microclimate changes due to alteration of plant cover, soil, water movement, and temperature patterns could conceivably affect herbaceous plant communities of vernal pools. Destruction of cyanobacterial associations may also permanently change composition and productivity in certain pool zones and negatively affect the loss of pool-edge moisture status and associated species. Pools altered by hydrologic and other related factors, i.e., nutrient status, are often invaded by exotic taxa and dominated by relatively large populations of only a few native species. Analysis of vernal pools might assume that habitat quality is positively correlated with species density; this could be misleading for rare species conservation since density alone may not be positively correlated with habitat quality (Van Horne 1983). Conservation issues involving successional seres of vernal pools are also relatively unexplored. Zedler (1987) considered it a mistake to assume that pools and non-pools are the only category of vegetation in vernal pool landscapes. If late successional or 312 PHY T@GEO GDA Apnil 1996 volume 80(4):296-327 disturbed silt-pools do support a dormant seed stock of obligate vernal pool species which only rarely germinate except under optimal hydrologic conditions, then these habitats should also be identified and protected, perhaps as a subclass of vernal pools. During drought years in southern coastal Califomia, Plantago elongata Pursh may be an indicator of silted, drying liquefaction pools. Twisselmann (1967) indicated that vernal pools in Kern County typically exist for years without water and/or the ce of vernal pool vegetation. Apparently, there is a continuum of water duration characteristics from which vernal pools in the strict sense and other ephemeral wetlands within vernal pool terrain develop (Zedler 1987). Other organisms, including fairy shrimp, also occupy the full range of intermittent pool types in vernal pool landscapes (Brown, ef al. 1993). From a conservation perspective, the most practical approach is ecosystem identification proposed by Sawyer & Keeler-Wolf (1995) rather than the simplified vascular plant community classification. Species found in vernal pools that are more abundant outside pool habitat in surface water slope wetlands in coastal sage scrub, i.e., Eryngium aff. vaseyi J. Coulter & Rose and Trifolium variegatum Nutt., may indicate the importance of conserving small ephemeral wetlands or seepage habitats in liquefaction-type terrain. A mosaic of pool complexes incorporating distinct edaphic types and serial stages is suggested. Cyanobacteria may play an important role in promoting maximum biodiversity in southern California’s vernal pools. Microcoleus and/or pools designed to duplicate the recurrent flux hypothesis should be incorporated into habitat mitigation plans since artificially created pools frequently lack species abundance and cover comparable to natural pools of the same region (Ferren & Gevirtz 1990; Zedler, et al. 1993). THE FLORA This preliminary compendium of cryptogamic and phanerogamic flora is the result of surveys conducted between the spring of 1993 and the spring of 1996 at San Clemente and San Onofre State Beaches, and infrequent visits to other Orange County sites listed below. Most noteworthy is the varied composition associated with duripan or claypan soils, and cycles of abundance which vary from year to year. Several of the species recorded during this study appeared only during the unusually wet spring of 1993 (1l-inches above-average precipitation; Orange County Environmental Management Agency 1996), possibly due to replenished inundation and germination regimes; other species absent or rare during wet years were recorded only during subsequent dner periods. Rare or regionally uncommon vascular plants of Orange or San Diego counties collected during this study include: Alopecurus saccatus Vasey, Atriplex pacifica Nelson, Brodiaea jolonensis Eastw., Crassula aquatica (L.) Schonl., Deschampsia danthonioides (Trin.) Benth., Dudleya blochmaniae (Eastw.) Moran, Eryngium aff. vaseyi, Harpogonella palmeri A. Gray Hordeum intercedens Nevski, Microseris douglasii (DC.) Schultz-Bip. subsp. platycarpha (A. Gray) Chambers, Marsilea vestita Hook. & Grev., Muilla maritima (Torrey) S. Watson, Myosurus minimus L., Navarretia prostrata A. Gray, Pilularia americana A. Braun, Psilocarphus brevissimus, Psilocarphus tenellus Nutt., and Senecio aphanactis E. Greene. Riefner & Pryor: Southern California vernal pools 313 Voucher specimens have been deposited in the University of California, Irvine, Museum of Systematic Biology (IRVC) and selected duplicates have been placed in the herbarium of the Rancho Santa Ana Botanic Garden (RSA). LEGEND * Indicates a specialist which is generally restricted to vernal pools or an introduced taxon (!*) characteristic of the vernal pool community in coastal California according to Zedler (1987). + Indicates an aquatic generalist that is more common in other aquatic, marsh, or seepage habitats. ! Indicates a non-native species that may tolerate inundation/saturation. o Indicates a native species occurring in diverse habitats that can tolerate limited periods of saturation/inundation. 4 Indicates a native or exotic species that may grow near vernal pools but does not grow in the saturation zone and cannot tolerate extended inundation. + Indicates extreme fluctuations in the population size/vigor or the presence/absence of a species recorded during the study period which may be attributed to variation in environmental conditions. R_ Rare at the site. I Occasional to infrequent. C Common. S BSurf Beach Unit, San Onofre State Beach, San Diego Dunpan Vemal Pools; Location: San Diego County. San Onofre State Beach. Surf Beach Unit (San Onofre USGS 7.5' Quadrangle, T9S, R7W, Section 24); Soils: Soils are classified as Carlsbad gravelly loamy sand, which is gently sloping and characterized by an iron-silica cemented duripan (Soil Conservation Service 1973). T Trestles Unit, San Onofre State Beach, San Diego Claypan Vemal Pools and Swales; Location: San Diego County. San Onofre State Beach. Trestles Natural Preserve Unit (San Clemente USGS 7.5’ Quadrangle T9S, R7W, Section 14); Soils: Visalia Series are mapped for this region; these have moderately rapid permeability and support soil inclusions such as Placentia soils which contain a sandy clay subsoil of very slow permeability (Soil Conservation Service 1973). SCSan Clemente State Beach, Orange County Liquefaction-Origin Vernal Pools; Location: Orange County. San Clemente State Beach (San Clemente USGS 7.5’ Quadrangle T9S, R7W, Section 10); Soils: Soils are classified as Myford Series, very slowly permeable, moderately alkaline, with a clay-nch subsoil (Soil Conservation Service 1978). O Additional Orange County Pools-Locations: Rancho Laguna-Laguna Beach (Laguna Beach USGS 7.5’ Quadrangle T8S, R8W, Section 31), Dana Hills (Dana Point USGS 7.5’ Quadrangle T8S, R8W, Section10/15), and San Clemente (San Clemente USGS 7.5’ Quadrangle T8S, R7W, Section 32); Soils: Myford sandy loam, rarely the moderately slowly permeable Botella clay loam (Soil Conservation Service 1978). 314 PHYTOLOGIA April 1996 volume 80(4):296-327 PRELIMINARY INVENTORY OF SPECIES INHABITING VERNAL POOLS AND ADJACENT UPLAND TERRAIN CRYPTOGAMIC PLANTS CYANOBACTERIA: o Microcoleus vaginatus (Vauch.) Gom., filamentous cyanobactenum + Nostoc sp., heterocystous cyanobacterium = fe QD BRYOPHYTES: o Asterella sp., liverwort - Rt - “ Bryum argenteum Hedw., moss R Cc I o B. bicolor Hedw., moss - R R o B. gemmiparum De Not., moss - - o B. pseudotriquetrum (Hedw.) Gaertn., Meyer & Schreb., moss - “ Ceratodon sp., moss - 0 Ceratodon purpureus (Hedw.) Bnid., moss Claopodium wippleanum (Sull.) Ren. & Card., moss - - Nf o Fossombria longiseta Aust., liverwort - I - o Funaria hygrometrica Hedw., cord moss R C R Riccia glauca L., liverwort - It R. nigrella DC., liverwort - R R. trichocarpa M.A. Howe, liverwort - R R ce aa’,”) ' WDA Scleropodium tourettei (Brid.) L. Koch, moss - Timmiella crassinervis (Hampe) L. Koch, moss - Tortula ruralis (Hedw.) Gaertn., Meyer & Schreb., moss : - R Weisia controversa Hedw., moss - - - >>+ >00 A toa ap Dee oo eae > Ayn LICHENS: (Terricolous species only; those not marked cyanolichen have green photobionts.) Acarospora cf. schleicheri (Ach.) A. Massal. - - - Caloplaca sp. - - Catapyrenium lachneum (Ach.) R. Sant. R : Cladonia sp. (sterile), pyxie cups - : Cladonia furcata (Hudson) Schrader - R - Cladonia scabriuscula (Delise) Ny). - 7 R Collema sp., cyanolichen - R Collema texanum Tuck., cyanolichen R - Diploschistes actinostomus (Ach.) Zahlbr. . - Lecanora cf. argopholis (Ach.) Ach. - - - Leprocaulon microscopicum ( Vill.) Gams ex D. Hawksw. - - I Leproloma sp. - - - > SSO Or > > > © eet Sos am wz > Riefner & Pryor: Southern California vernal pools A Psora decipens (Hedw.) Hoffm. - R A Rinodina bolanderi H. Magn. - - A Trapeliopstis sp. - : PHANEROGAMIC PLANTS FERNS: Marsileaceae, marsilea family * Marsilea vestita Hook. & Grev. subsp. vestita, clover fern (RSA) - - * Pilularia americana A. Braun, pillwort(RSA) Cf - MONOCOTS: Cyperaceae, sedge family + Eleocharis macrostachya Britton, pale spike-rush - - Iridaceae, iris family o Sisyrinchium bellum S. Watson, blue-eyed grass | - Juncaceae, rush family + Juncus bufonius L. var. bufonius, toad rush (RSA) = Ij o J. mexicanus Willd., Mexican rush - . Liliaceae, lily family A Bloomeria crocea (Torrey) Cov., common golden star - R o Brodiaea jolonensis Eastw., mesa brodiaea (RSA) - - ® Calochortus splendens Benth., splendid mariposa lily - - A Chlorogalum parviflorum S. Watson, soap plant - - A Dichelostemma capitatum Alph. Wood, blue dicks R : 0 Muilla maritima (Torrey) S. Watson, common muilla (RSA) : - Poaceae, grass family * Alopecurus saccatus Vasey, foxtail grass (RSA) . !+Agrostis viridis Gouan, bent grass - “ Avena barbata Link, slender wild oat < ! Avena fatua L., wild oat I !+ Briza minor L., little quaking grass Rt “ Bromus diandrus Roth, ripgut grass I ! Bromus hordeaceus L., soft chess S “ Bromus madritensis L. subsp. rubens (L.) Husnot, foxtail chess Cc I !+Crypsis schoenoides (L.) Lam., swamp grass_- - !+Cynodon dactylon (L.) Pers., bermuda grass R : * Deschampsia danthonioides (Trin.) Benth., annual hairgrass (RSA) - RT 0 Distichlis spicata (L.) E. Greene, salt grass R - ! Gastridium ventricosum (Gouan) Schinz & Thell., nit grass (RSA) - : Ct | o> hapa da pall 2 mal _ ‘e ' AAD yA DW = — 315 316 PHYTOLOGIA Apnil 1996 !4 Hainardia cylindrica (Will.) Greuter (RSA) o Hordeum intercedens Nevski, barley (RSA) A Hordeum murinum L. subsp. leporinum (Link) Arcang., barley A Lamarckia aurea (L.) Moench, goldentop ! Lolium multiflorum Lam, Italian ryegrass !+ Lolium perenne L., perennial ryegrass A Nassella lepida (A. Hitchc.) Barkworth, foothill needle- A N. pulchra (A. Hitchce.) Barkworth, purple needle-grass ! Phalaris minor Retz., littleseed canary grass (RSA) !* Phalaris paradoxa L., paradox canary grass ! Poa annua L., annual bluegrass !+ Polypogon monspeliensis (L.) Desf., rabbit-foot grass ! Vulpia myuros (L.) C. Gmelin var. myuros, rattail fescue A V. myuros var. hirsuta (Hackel) Asch. & Graebner, rattail fescue DICOTS: Amaranthaceae, amaranth family ! Amaranthus albus L., tumbleweed o A. blitoides S. Watson, prostrate amaranth \ A. deflexus L., low pigweed Anacardiaceae, sumac family “ Rhus integrifolia (Nutt.) Brewer & S. Watson, lemonadeberry Aizoaceae, fig-marigold family ® Malephora crocea (Jacq.) Schwantes, croceum iceplant “ Mesembryanthemum nodiflorum L., slender-leaved iceplant Apiaceae, celery family * Eryngium aff. vaseyi J. Coulter & Rose, coyote-thistle (RSA) ! Foeniculum vulgare Miller, fennel Asteraceae, sunflower family o Amblyopapus pusillus Hook. & Armm., coast weed o Ambrosia psilostachya DC., western ragweed 0 Baccharis pilularis DC., coyote brush + B. salicifolia (Ruiz Lopez & Pavon) Pers., mule fat “ Centaurea melitensis L., yellow star-thistle ! Chamomilla suaveolens (Pursh) Rydb., pineapple weed “ Chrysanthemum coronarium L., crown daisy 0 Conyza canadensis (L.) Crong., horseweed If c — R I S yee P| volume 80(4):296-327 —s Ct ee CO ee ae — AAD — i 1 3 a NAW A BW! v2) ' Riefner & Pryor: o Conyza coulteri A. Gray, Coulter’s horseweed_ - !+ Cotula coronopifolia L., brass-buttons Cc + Euthamia occidentalis Nutt., western goldenrod - A Filago californica Nutt., fluffweed - A F. gallica L., narrow-leaved filago - ! Gnaphalium luteo-album L., weedy cudweed * Gnaphalium palustre Nutt., lowland cudweed = Rf © Grindelia camporum E. Greene var. camporum, gumplant - “ Hedypnois cretica (L.) Dum.-Cours., Crete hedypnois - o Hemizonia fasciculata (DC.) Torrey _&A. Gray, tarweed 6 o Hemizonia paniculata A. Gray, San Diego tarweed - ! Hypochaeris glabra L., cat’s ear . o Isocoma menziesii (Hook. & Am.) G. Nesom var. menziesii, goldenbush + I. menziesii var. vernioides (Nutt.) G. Nesom, goldenbush (RSA) - ! Lactuca serriola L., prickly lettuce I 0 Lasthenia californica Lindley, common goldfields - o Layia platyglossa (Fischer & C. Meyer) A. Gray, tidy-tips - 0 Micropus californicus Fischer & C. Meyer, slender cottonweed - 0 Microseris douglasii (DC.) Schultz-Bip. subsp. platycarpha (A. Gray) Chambers, small-flowered microseris (RSA) & Pluchea odorata (L.) Cass., salt marsh fleabane_ - Psilocarphus brevissimus Nutt. var. brevissimus, woolly-heads (RSA) ie * P. tenellus Nutt. var. tenellus, slender woolly-heads (RSA) - Senecio aphanactis E. Greene, rayless ragwort (RSA) ! Senecio vulgaris L., common groundsel ! Sonchus asper (L.) Hill subsp. asper, prickly sow-thistle I ! Sonchus oleraceus L., common sow-thistle 0 Stebbinsoseris heterocarpa (Nutt.) Chambers, derived microseris R “ Uropappus lindleyi (DC.) Nutt., silver puffs . Boraginaceae, borage family ® Cryptantha micromeres (A. Gray) E. Greene, minute-flowered cryptantha (RSA) - © Harpagonella palmeri A. Gray, Palmer’s grappling hook - * Plagiobothrys acanthocarpus (Piper) I.M. Johnston, adobe popcorn flower (RSA) Cc * + (eo) ‘ e ! — Southern California vernal pools | I I G¢ | ole of ee | Rf aa 2 — v2) w I aa ay ibe | ON oo fold Rf Rf = ~— ae 317 318 PHYTOLOGIA Apnmil 1996 A P. collinus (Philbr.) I.M. Johnston var. fulvescens (1.M. Johnston) Higgins, rough popcorn flower - o P. nothofulvus (A. Gray) A. Gray, popcorn flower - Brassicaceae, mustard family ! Brassica nigra (L.) Koch, black mustard - © Cardamine californica (Torrey & A. Gray) E. Greene var. integrifolia (Torrey & A. Gray) Rollins, milkmaids - o Lepidium nitidum Torrey & A. Gray, shining peppergrass Cc Callitrichaceae, starwort family * Callitriche marginata Torrey, wallow starwort Rf Caryophyllaceae, pink family “ Silene gallica L., common catchfly ! Spergularia bocconii (Scheele) Merino, sand-spurrey R 0 Spergularia macrotheca (Hornem.) Heynh. var. macrotheca, sticky sand-spurrey (RSA) - ! Spergularia villosa (Pers.) Cambess., villous sand-spurrey ] Chenopodiaceae, goosefoot family “ Atriplex lindleyi DC., a saltbush (RSA) R 0 A. pacifica Nelson, south coast saltbush (RSA) - ! A. rosea L., tumbling oracle - ! A. semibaccata R. Br., Australian saltbush & o A. serenana Nelson, bractscale (RSA) - ! Chenopodium album L., |amb's quarters - ! C. ambrosioides L., Mexican tea - ! Salsola tragus L., tumbleweed R Convolvulaceae, morning glory family 0 Cressa truxillensis Kunth, alkali weed (RSA) - ! Convolvulus arvensis L., bind weed R Crassulaceae, stonecrop family * Crassula aquatica (L.) Schonl., pygmy stonecrop (RSA) CT o C. connata (Ruiz Lopez & Pavon) A. Berger, pygmy-weed Cc ‘ Dudleya blochmaniae (Eastw.) Moran subsp. blochmaniae, Blochman’s dudleya - “ D. edulis (Nutt.) Moran, ladies-fingers - Elatinaceae, waterwort family * Elatine brachysperma A. Gray, waterwort (RSA) It Euphorbiaceae, spurge family ! Chamaesyce maculata (L.) Small, spotted spurge (RSA) I Fabaceae, pea family 0 Trifolium depauperatum Desv. var. amplectens (Torrey & A. Gray) L.F. McDermott, pale sack clover (RSA) Cc eee Pe SC ee —- ye) volume 80(4):296-327 Riefner & Pryor: o T. variegatum Nutt., white-tip clover (RSA) Geraniaceae, geranium family ! Erodium botrys (Cav.) Bertol., broadleaf filaree (RSA) A E. cicutarium (L.) L’Hér., red-stemmed filaree A E. moschatum (L.) L’Hér., white-stemmed filaree Lamiaceae, mint family ! Marrubium vulgare L., horehound Lythraceae, loosestrife family !* Tythrum hyssopifolia L., \oosestrife Malvaceae, mallow family ! Malva parviflora L., cheeseweed 0 Malvella leprosa (Ortega) Krapov., alkali-mallow Molluginaceae, carpet-weed family !+Glinus lotoides L., carpet-weed Oxalidaceae, oxalis family A Oxalis pes-caprae L., Bermuda buttercup Plantaginaceae, plantain family o Plantago elongata Pursh, alkali plantain (RSA) A P. erecta E. Morris, California plantain ! P. virginica L., plantain (RSA) Polemoniaceae, phlox family A Linanthus dianthiflorus (Benth.) E. Greene, ground-pink. o Navarretia atractyloides (Benth.) Hook. & Am., holly-leaved skunkweed (RSA) * N. prostrata A. Gray, navarretia (RSA) Polygonaceae, buckwheat family ! Polygonum arenastrum Boreau, common knotweed !+Rumex crispus L., curly dock Portulacaceae, purslane family o Calandrinia ciliata (Ruiz Lopez & Pavon) DC., red maids Primulaceae, primrose family !* Anagallis arvensis L., scarlet pimpemel Ranunculaceae, buttercup family * Myosurus minimus L., little mousetail (RSA) 0 Ranunculus californicus Benth., California buttercup Rubiaceae, madder family o Galium aparine L., goose grass Scrophulariaceae, figwort family 0 Castilleja exserta (A.A. Heller) Chuang & Heckard, purple owl’s clover 4 Linaria canadensis (L.) Dum.-Cours., blue toadflax + Veronica peregrina L. subsp. xalapensis (Kunth) Pennell, purslane speedwell (RSA) or,” Southern California vernal pools i aw 7, aa as aw 320 PHY TOLCGIA Apnil 1996 volume 80(4):296-327 ACKNOWLEDGMENTS We are deeply indebted to the following individuals for their indispensable support and contributions to this project: Dr. Roy Shlemon provided geologic interpretations; Dr. Gerald Kuhn provided aerial photographs and interpretation of liquefaction issues; Steve Boyd, Rancho Santa Ana Botanic Garden, annotated vascular plants; Dr. Jayne Belnap annotated cyanobacteria and provided helpful discussion on the ecology of Microcoleus, Dr. Peter Bowler, University of California, Irvine, shared his knowledge of Orange County vernal pools and provided herbarium and laboratory space; David Bramlet, Santa Ana, California, identified Hordeum intercedens and provided helpful discussion; Dr. George Cox, San Diego State University, provided information that assisted in the preliminary identification of mima mounds; and Dr. William Bretz, University of California Natural Reserve System, provided aenal photographs of Orange County. 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Studies from the Herbarium, No. 8, California State University, Chico, California. , C.K. Frazier, & C. Black. 1993. Habitat creation as a strategy in ecosystem preservation: an example from vernal pools in San Diego County. Jn: J. Keeley (ed.). Interface Between Ecology and Land Development in California. Southern California Academy of Sciences, Los Angeles, California. NOW AVAILABLE, PHYTOLOGIA MEMOIRS, Volume 10 THE COMPS OF MEXICO, Volume 6 TAGETEAE AND ANTHEMIDEAE B.L. TURNER 1996, 8 1/4 x 10 3/4 in., 97 pp., $29.95, ISBN 0-9642085- 1-2. The first of an anticipated ten volume series on the Asteraceae of México, this volume includes the tribes Tageteae and Anthemideae. Later 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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