om and j ecological publica tion oe 1998 Rei ace Dea Te geal - QUIROZ.G. “Dail, R PALACIOS C., “aM. ‘ARREGUIN S., Morfologia de pea ~~. Jos granos de polen de la familia Polygonaceae del Valle de México, México. 1” a WEBER, W.A., New names and combinations in Asteraceae: Heliantheae - | Pee esCliiiae i.e a ota: es tele Stee ee Yael ig Se ccar, Citi ag gs |i: aes _ MACROBERTS, M. H & BR. MACROBERTS, Welland status of hillside agen SOBs wetland Pine savannas, and mick bogs in the West Gulf Coastal Plain. Meas acs et cipeyhr ae taneiek ene acon ta Bipimern prog ek eae ve beds eae nee ewes Meee ai Dare MACROBERTS, M. H. & B.R. MACROBERTS, Community classification of — -. West Gulf Coastal Plain bog communities: a floristic assessment. ......... Wd MACROBERTS B.R. & M.H. MACROBERTS, Floristics of wetland pine - savannas in the Big Thicket National Preserve, southeast Texas. ........... 40 = NEY LAND, R., H.A. MEYER, & H. HARRINGTON, Woody vegetation of _.~ Aongleaf pine communities in Calcasieu Parish, Louisiana..................:. Shee _*MACROBERTS, B:R.. a M.H. MACROBERTS, Floristics of muck bogs. in east central Texas. ....... bien ay Seek alttty cis rice aun eee ueM Ucn Res ge Ge Gr ~ PERLMUTTER, G.B., Sex morph descriptions of ‘Malosna laurina "=. (Anacardiaceae), a polygamodioecious SPECIES Sika ke hee ete as gat PORGOKS-TECOIVEU: oer. cco ici hae caw titi reeds oes Rs ay TAG uae Bante a 8 _ Publication dates, volume 84, .2.00.......0...20.05 ORS EN ae Pome Smee asf 81 _ Index to reviewers, volume 84. ............2...... EE OTD AAMC ERR Cnt. Pai ~ Index to authors, VONMIMEG B4 25 os Ales tikes oan fe hoy Sn RON Se Sie eee Tat 83 pew names in this i issue of Phytologia. Re caavea vee. erat LAN RCS nama dao oh Oe LIBRARY NOV =9 (9. NEWYORK BOTANICA! CAD. xP ot] ae: a s - PHYTOLOGIA “An international journal to expedite plant systematic, Phytogeographicel | and: feb ogiea! sepieds tion SVL: Ba = : x July 1998 Hae: re No. 1 oon = - QUIROZ 5 DL R. PALACIOS C., -& M. ARREGUIN S., Morfologia de § --los granos de polen de la familia Polygonaceae del Valle de México, México. 1 ~ we Be. WEBER, W.A., New names and combinations i in Asteraceae: Heliantheae - Bete ESCH A. bs acer eg oBase ewes Cases a vue ea yen yar ob Sou eeaig ener Make Agi eS i ages ~. MACROBERTS, M.H. & B.R.. ‘MACROBERTS, Wetland status of hillside x ga, wetland pine savannas, and muck bogs in the West Gulf Coastal Plain. Be Gi es ciety es Lan wk oak ema ta Pip nen prod Gos eae Uns Me eee te eats 22 ~ MACROBERTS, M.H. & B.R. MACROBERTS, Community classification of '-. West Gulf Coastal Plain bog communities: a floristic assessment. ......... ao: *- MACROBERTS B.R. & M.H. MACROBERTS, Floristics of wetland pine -. savannas in the Big Thicket National Preserve, southeast Texas. ........... 40 = a NEYLAND, R., H.A. MEYER, & H. HARRINGTON, Woody vegetation of ~~ Jongleaf pine communities in Calcasieu Parish, Louisiana..................:. Sle __ MACROBERTS, B-R. & M.H. MACROBERTS, Flonistics of muck bogs. in ve CASU-CONMAL TERAS ee oars es en lain es wea ee eae ema eee Eke Sas 6s: - PERLMUTTER, G.B., Sex morph ‘descriptions of Malosma Jaurina ae (Anacardiaceae), a polygamodicecious Species. ....... Reeeneess ates reseee ‘Tar Mer S TECOIVEU: 62 cine ect pca so ese tan Pats Re bewes aes SNe ghia Citi Sate nee ~ Publication dates; volume 84. ....0..0.c..00. cee. Re ts oe ea eR Ia 81 = Index to reviewers, volume 84. .......5.....c50..0cuesedeeee ona Sak At ote 82 Btndex (0 authors, Voluine 84.24 5.2.. 4 pthc se or iad clas api cae rat 83 - New names in this issue of Phytologia. Baca eres ees Savece mre 84 ae Nov = 9 0” co o os NEW YORK : | - ROTANICAL CARD. I RS ENS RE ee os ytd roa ae «ees Meth a ae. <, e, Phytologia (July 1998) 85(1):1-18. MORFOLOGIA DE LOS GRANOS DE POLEN DE LA FAMILIA POLYGONACEAE DEL VALLE DE MEXICO, MEXICO’ David Leonor Quiroz Garcia”, Rodolfo Palacios Chavez’, & Marfa de la Luz Arreguin Sanchez? Departamento de Botanica, Escuela Nacional de Ciencias Bioldgicas, I.P.N., Plan de Ayala y Carpio, Col. Santo Tomas 11340 México, D.F., MEXICO ABSTRACT Pollen descriptions of three genera and eighteen species of Polygonaceae from Valle de México are studied and illustrated. The taxa are: Eriogonum jamesii Benth., Polygonum amphibium L., P. argyrocoleon Steud., P. aviculare L., P. coccineum Muhl., P. convolvulus L., P. hartwrightii A. Gray, P. hydropiperoides Michx., P. lapathifolium L., P. mexicanum Small, P. punctatum Ell., Rumex acetosella L., R. conglomeratus Murr., R. crispus L., R. flexicaulis Rech., R. mexicanus Meisn., R. obtusifolius L., and R. pulcher L. Observations of pollen grains were carned out with scanning electron microscopy and light microscopy. Pollen is usually spheroidal, subprolate, suboblate; tricolporate, tetracolporate, pericolpate, and the omamentation fossulate, foveolate, microreticulate, reticulate, microverrucate, and microechinate. KEY WORDS: Palynology, Polygonaceae, Valle de México, México RESUMEN Se estudia e ilustra la morfologia polinica de los tres géneros y las diesiocho especies de la familia Polygonaceae del Valle de México: Eriogonum jamesii Benth., Polygonum amphibium L., P. argyrocoleon Steud., P. aviculare L., P. coccineum Muhl., P. convolvulus L., P. hartwrightii A. Gray, ‘Proyecto apoyado por la Direccién de Estudios Profesionales e Investigacién del Instituto Politécnico Nacional. *Becarios de la Comisién de Operaciones y Fomento Académico. l 2 PHYTOLOGIA July 1998 volume 85(1):1-18 P. hydropiperoides Michx., P. lapathifolium L., P. mexicanum Small, P. punctatum Ell., Rumex acetosella L., R. conglomeratus Murt., R. crispus L., R. flexicaulis Rech., R. mexicanus Meisn., R. obtusifolius L., y R. pulcher L. Las observaciones de los granos de polen se realizaron con microscopio de luz y microscopio electrénico de barrido. El polen se observé esferoidal, subprolato, suboblato; tricolporado, tetracolporado, pericolpado o periporado; la ornamentacién es fosulada, foveolada, microrreticulada, reticulada, microverrugada, y microequinada. PALABRAS CLAVE: Palinologfa, Polygonaceae, Valle de México, México INTRODUCCION La familia Polygonaceae estd constituida por 40 géneros y cerca de 800 especies (Lawrence 1969). Su distribucién es cosmopolita, se le encuentra principalmente en las regiones templadas del hemisferio norte. En el Valle de México prosperan tres géneros, diesiocho especies, y dos subespecies (Rzedowski & Rzedowski 1979). Melchior (1964), divide a la familia en tres subfamilias: | Enogonoideae, Polygonoideae, y Coccoloboideae cada una con dos tribus. Los taxa que se incluyen en este trabajo se ubican en las dos primeras subfamilias. ANTECEDENTES La morfologia del polen de la familia Polygonaceae ha sido estudiada por autores como Wodehouse (1931) que incluye las descripciones de once géneros y 27 especies, elabora una clave y hace comentarios sobre la evolucién de los tipos polinicos. Erdtman (1952) considera el polen de diesiocho géneros y 170 especies, sefialando las caracteristicas morfoldgicas de las especies que ilustran la estratificacién de la exina. Kapp (1969) incluye en sus claves polinicas tres géneros y tres especies. Huang (1972) describe de Taiwan el polen de dos géneros y 27 especies de esta familia. Markgraf & D“Antoni (1978) estudian el polen de tres géneros y tres especies de Argentina. Nowicke & Skvarla (1979) sefialan que las poligondceas tienen una gran diversidad de tipos polinicos debido a la variacién en todas sus caracteristicas como son: aperturas, ornamentacién y estructura de la exina. van Leeuwen et al. (1988) examinan el polen de 47 especies de la familia y las separan en diez tipos polinicos. Roubik & Moreno (1991) describen el polen de tres géneros y ocho especies de la isla de Barro Colorado, Panama. Palacios-Chavez et al. (1991) incluyen de la reserva de Sian Ka’an el polen de cinco especies. Quiroz-Garcia et al. (1995) describen el polen de cuatro géneros y ocho especies del estado de Jalisco, que prosperan en clima cdlido. Quiroz G. etal.: Pollen of Polygonaceae 3 METODOLOGIA El polen fue tomado de ejemplares depositados en el herbario ENCB de la Escuela Nacional de Ciencias Biolédgicas. Cuando no se pudo tomar polen de esa coleccién, por carecer de flores, el material se obtuvo de otros ejemplares del pafs, previa corroboracién de la identificaci6n de los mismos. Las muestras se dividieron en dos partes; una fue tratada con la técnica de acetdlisis de Erdtman (1943) para observaciones al microscopio de luz (ML) y otra parte, sin tratamiento quimico, se utiliz6 para el microscopio electrénico de barrido (MEB). El polen fue recubierto con oro, para observaciones en un microscopio JEOL-JSM 35. La terminologfa empleada es la propuesta por Punt et al. (1994). DESCRIPCION DE LOS GRANOS DE POLEN Eriogonum jamesii Benth. Mesa del Cinco, Sierra Madera del Carmen, Mpio. Villa de Ocampo, Coahuila. M.F. Robert y J. Passini 5522 (ENCB). Lamina 1, Figuras 1-6. Polen tricolporado, tectado, subprolato de 49(57)61 x 36(43)51 ym. P/E=1.3. Vista polar subangular de 36(41)44 ym de didmetro. Exina de 3.8 um de grosor, nexina de menor espesor que la sexina; al ML y MEB foveolada, los foveolos se observan en grupos. Colpos largos y delgados. Poros lolongados de 5(7)8 um de largo x 4(5)6 um de ancho. Area polar 0.26, mediana. Polygonum amphibium L. Sierra de Judrez, ca. de Laguna Hanson, Baja California. R. Moran 2919] (ENCB). LaminalI, Figuras 7-10. Polen pericolpado, semitectado, esférico de 53(59)65 ym de didmetro. Exina de 4 yum de grosor, con la nexina de menor espesor que la sexina, al ML y MEB reticulada. Colpos de 15 a 27, orentados como los lados de pentdgonos regulares, de 7.0(8.5)10.0 um de largo x 1.1 um de ancho. Polygonum argyrocoleon Steud. Sn. Gregorio Atlapulco, Distrito Federal. F. Pdez s/n, 26-X]-72 (ENCB). Laminal, Figuras 11-14 y Laminall, Figuras 15 y 16. Polen tncolporado a tetracolporado, zonorado, tectado, subprolato de 26(28)30 x 19(21)22 um. P/E=1.28. Vista polar circular de 19(21)24 wm de didmetro. Exina de 2.0 wm de grosor, con la nexina de menor espesor que la sexina; al ML y MEB microverrugado. Colpos largos y delgados cubiertos con membranas 4 PHYTOLOGIA July 1998 volume 85(1):1-18 microverrugadas. Colpos transversos que se unen a la altura del ecuador, formando una banda continua de 2.6 ym de ancho. Indice del drea polar 0.38, media. Polygonum aviculare L. Terrenos de Jaltepec, Mpio. de Ajapulco, Estado de México. A. Ventura 2148 (ENCB). Lamina II, Figuras 17-22. Polen tricolporado, tectado, subprolato de 33(37)41 x 27(29)30 um. P/E=1.28. Vista polar semiangular de 27.0(29.6)33.0 um de didmetro. Exina de 2.0 um de grosor, con la nexina de menor espesor que la sexina; al ML y MEB microequinada. Colpos largos y delgados. Polen zonorado, colpos transversales que se unen a la altura del ecuador, formando una banda continua de 3.3 um de ancho. Indice del drea polar 0.35, media. Polygonum coccineum Muhl. Presa de Cuevecillas, 5S Km al SW de Huehuetoca, Estado de México. A. Herndndez 50 (ENCB). Lamina II, Figuras 23-27. Polen pericolpado, semitectado, esférico de 48(56)65 um de didmetro. Exina de 3.8 ym de grosor, con la nexina de menor espesor que la sexina; al ML y MEB reticulada. Colpos 27 a 30, onentados como los lados de un pentdgono regular, de 6(7)9 um de largo x 0.6(1.0)1.5 um de ancho. Polygonum convolvulus L. Cerro Penférico, ca. del Hospital del Nifio, Mpio. Saltillo, Coahuila. R. Ldpez 751 y J. A. Villarreal (ENCB). Laémina III, Figuras 28-34. Polen tricolporado, tectado, subprolato de 26(27)30 x 20(22)24 um. P/E= 1.22. Vista polar semiangular de 20(21)22 wm de didmetro. Exina de 1.3 a 2.1 wm de grosor con la nexina de menor espesor que la sexina, ornamentacién al ML reticulada, al MEB microequinada. Colpos delgados con terminaciones agudas. Polen zonorado, colpos transversos que se unen a la altura del ecuador, formando una banda continua de 2.8 um de ancho. Indice del drea polar 0.42, media. Polygonum hartwrightii A. Gray. Xochimilco, orilla del canal, Distrito Federal. J. Rzedowski 28143 (ENCB). LaminallI, Figuras 35-38. Polen pericolpado, semitectado, esférico de 57(67)72 um de didmetro. Exina de 4.2 um de grosor, con la nexina de menor espesor que la sexina, superficie al ML y MEB reticulada, con verrugas en los Iimenes. Colpos 27 a 30, orientados como los lados de un pentdgono regular, de 8.0(9.3)12.0 jum de largo x 1.2 jum de ancho. Quiroz G. etal.: Pollen of Polygonaceae 5 LOK. $3208 1] 12 13 14 LAMINA | Liminal. Eriogonum jamesii. 1. Vista ecuatorial mostrando el grosor de la exina. 2. Vista ecuatorial superficial. 3. Vista polar. 4. Vista ecuatonal al MEB. 5S. Acercamiento superficial al MEB. 6. Vista polar al MEB. Polygonum amphibiwm. 7. Seccidn Optica. 8. Vista superficial. 9. Vista general del grano de polenal MEB. 10. Detallede la ornamentacion al MEB. Polygonum argyrocoleon. 11. Vista ecuatorial, secci6n Optica. 12. Vista ecuatonal superficial. 13. Vista polar seccién 6ptica. 14. Vista polar supeificial. Barra= 10 um. PREY FOLOGLA July 1998 ; - FOR BOOB « LAMINA Hi Laminall. Polygonum argyrocoleon. 15. Vista total del grano de polen al MEB. 16. Acercamiento superficial al MEB. Polygonum aviculare. 17. Vista ecuatorial, secci6n Optica. 18. Vista ecuatorial superficial. 19. Vista polar, seccién Optica. 20. Vista polar superficial. 21. Vista ecuatorial al MEB. 22. Vista polar al MEB. Polygonum coccineum. 23. (Seccidn Optica. 24. Vista superficial. 25. Omamentaci6n. 26. Vista general del grano de polen al MEB. 27. Detalle de la Ornamentaci6n al MEB. Barra= 10 um Quiroz G. etal.: Pollen of Polygonaceae 40 LAMINA, Ul Laminalll. Polygonum convolvulus. 28. Vista ecuatorial, secci6n d6ptica. 29. Vista ecuatorial superficial. 30. Vista polar, secci6n 6ptica. 31. Vista polar superficial. 32. Vista total del grano de poien al MEB. 33. Acercamiento superficial al MEB. 34. Vista polar al MEB. Polygonum hartwrightii. 35. Vista total del grano de polen. 36. Detalle de la omamentacién. 37. Vista general al MEB. 38. Acercamiento superficial al MEB. Polygonum hydropiperoides. 39. Seccién Optica. 40. Vista superficial. Barra= 10 um. 8 PHYTOLOGIA July 1998 volume 85(1):1-18 83488 1722 PABEY ALISE 192°5 52 LAMINA PY LdminalV. Polygonum hydropiperoides. 41. Detallede la omamentacion. Polygonum lapathifolia. 42. Seccién dptica. 43. Vista general del grano de polen. 44. Vista superficial. 45. Vista total al MEB. 46. Acercamientosuperficial al MEB. Polygonum mexicanum. 47. Seccién 6ptica. 48. Vista superficial. 49. Vista general del grano de polen al MEB. 50. Detallede la ornamentacién al MEB. Polygonum punctatum. 51. Seccién 6ptica. 52. Detalle superficial. Barra= 10 pm. Quiroz G. etal.: Pollen of Polygonaceae 9 M5 GBB 2 LAMINA V¥ La4mina V. Polygonum punctatum. 53. Vista total del grano de polen al MEB. 54. Acercamiento superficial al MEB. Rumex acetosella. 55. Vista ecuatorial, seccién Optica. 56. Vista polar, seccidn 6ptica. 57. Vista polar superficial. 58. Vista general del grano de polen al MEB. 59. Vista polar al MEB. Rumex conglomeratus. 60. Vista ecuatorial superficial. 61. Vista polar seccién Optica. 62. Vista ecuatorial al MEB. 63. Acercamiento superficial al MEB. 64. Vista polar al MEB. Barra= 10 pm. 10 PHYTOLOGIA July 1998 volume 85(1):1-18 19. G0 JEOLS LAMINA Vi L4mina VI. Rumex crispus. 65. Vista ecuatorial, secci6n 6ptica. 66. Vista polar superficial. 67. Vista polar, seccién Optica. 68. Vista general del grano de polen al MEB. 69. Detallede la ornamentaci6n al MEB. 70. Vista polar al MEB. Rumex flexicaulis. 71. Vista polar superficial. 72. Vista polar, secci6n Optica. 73. Vista general del grano de polen al MEB. 74. Vista polar al MEB. 75. Detalle de la ornamentacion al MEB. Barra= 10 um. Quiroz G. etal.: Pollen of Polygonaceae LZ LAMINA VI Ldmina VII. Rumex mexicanus. 76. Vista ecuatonal, secciédn Optica. 77. Vista ecuatorial superficial. 78. Vista polar, seccién Optica. 79. Vista polar superficial. 80. Acercamiento superficial al MEB. 81. Vista general del grano de polen al MEB. Rumex obtusifolia. 8&2. Vista ecuatorial, secci6n Optica. 83. Vista ecuatonal superficial. 84. Vista polar superficial. 85. Vista ecuatorial al MEB. 86. Vista polar al MEB. 87. Detallede la ornamentacién al MEB. Barra= 10 pum. 12 PHY TOLOGTA July 1998 volume 85(1):1-18 LAMINA VU _Lémina VIII. Rumex pulcher. 88. Vista ecuatorial, seccién 6ptica. 89. Vista ecuatorial mostrando los colpos. 90. Vista ecuatorial superficial. 91. Vista polar, secci6n Gptica. 92. Vista polar mostrando cuatro aberturas. 93. Vista polar superficial. 94. Vista ecuatorialal MEB. 95. Acercamiento superficial al MEB. 96. Vista polar al MEB. Barra= 10 um. Quiroz G. etal.: Pollen of Polygonaceae 13 Polygonum hydropiperoides Michx. Lago de Patzcuaro, en el canal de Sn. Pedrito frente al muelle, Mpio. Patzcuaro, Michoacan. A. Lot (ENCB). Ldmina III, Figuras 39-40 y LaminalV, Figura 41. Polen periporado, semitectado, esférico de 40(44)47 um de didmetro. Exina de 3.9 um de grosor con la nexina de menor espesor que la sexina, ornamentacién al ML y MEB reticulada, con verrugas y gemas en los lumenes. Poros circulares de 2.6(3.9)5.2 um de didmetro. Polygonum lapathifolium L. Ex Hacienda de Xalpa, Mpio. Huehuetoca, Estado de México. R. Rojas 148 (ENCB). LaminalV, Figuras 42-46. Polen periporado, semitectado, esf€rico de 36(40)50 um de didmetro. Exina de 3.8 pm de grosor, con la nexina de menor espesor que la sexina, ornamentacién al ML y al MEB reticulada, con gemas en los lumenes. Poros circulares de 2.6(3.0)3.9 um de didmetro. Polygonum mexicanum Small. Laguna de Sn. Vicente, Mpio. de Tepeapulco, Hidalgo. A. Ventura 226 (ENCB). LdminalV, Figuras 47-50. Polen periporado, semitectado, esférico de 54(65)72 um de didmetro. Exina de 4.9 um de grosor, con la nexina de menor espesor que la sexina, ornamentacién al ML y MEB reticulada, con verrugas y gemas en los lumenes. Poros circulares de 2.6(3.6)3.9 um de didmetro. Polygonum punctatum Ell. 1 Km al NE de Tenango del Aire, Estado de México. A. Pineda 436 (ENCB). LdminalV, Figuras 51-52 y Lamina V, Figuras 53-54. Polen periporado, semitectado, esférico de 48(53)56 um de didmetro. Exina de 5.2 um de grosor, con la nexina y sexina del mismo espesor, ornamentacién al ML y MEB reticulada, con gemas en los lumenes. Poros circulares de 3.9(4.1)5.2 am de didmetro. Rumex acetosella L. Sta. Cecilia, Delegacién de Xochimilco, Distrito Federal. F. Ventura 2877 (ENCB). Lamina V, Figuras 55-59. Polen tricolporado, algunas veces tetracolporado, semitectado, esferoidal de 20(25)28 x 23.0(25.6)26.0 um. P/E=1. Vista polar circular de 22.0(24.7)28.0 um de didmetro. Exina de 1.3 wm de grosor, con la nexina de menor espesor que la sexina, Ormamentaci6n al ML y MEB microrreticulada. Sincolpado, colpos delgados a 14 PHYTOLOGIA July 1998 volume 85(1):1-18 veces se unen en los polos. Colpo transverso de 2.6(2.9)3.9 wm de largo x 2.6(4.4)5.1 pm de ancho. Indice del drea polar 0.35, media. Rumex conglomeratus Murr. 1 Km S de Tultitlén sobre el camino a Tlalnepantla, Estado de México. J. Rzedowski 28993 (ENCB). Lamina V, Figuras 60-64. Polen tricolporado, semitectado, esferoidal de 27(30)33 x 26(27)30 um. P/E=1.11. Vista polar circular de 22.0(30.3)34.0 wm de didmetro. Exina de 1.3 wm de grosor con la sexina del mismo espesor que la nexina, ornamentaci6n al ML microrreticulada, al MEB foveolada con microespinas. Colpos delgados en ocasiones se unen en uno de los polos (sincolpado). Colpos transversos de 3.9(4.8)6.5 um de largo x 2.6 wm de ancho. Indice del drea polar 0.35, media. Rumex crispus L: 2 Km al NW de Zumpango, ca. canal del desagiie, Estado de México. A. Pineda 711 (ENCB). Lamina V1, Figuras 65-70. Polen tricolporado, algunas veces parasincolporado, semitectado, suboblato de 24(30)33 x 31(34)37 wm. P/E=0.88. Vista polar circular de 30.0(32.5)36.0 wm de didmetro. Exinade 1.6 um de grosor con la nexina de menor espesor que la sexina, ornamentacién al ML microrreticulada, al MEB con microespinas sobre los muros. Colpos delgados en ocasiones se unen en uno de los polos. Poro circular de 2.6(4.4)5.2 um de largo x 3.9 wm de ancho. Indice del area polar 0.28, media. Rumex flexicaulis Rech. Pedregal del Pulpito, Mpio. de Juchitepec, Estado de México. A. Ventura 1318 (ENCB). Lamina VI, Figuras 71-75. Polen pericolpado, tetracolporado, en menor proporcién tricolporado, semitectado, esferoidal de 23(25)28 x 24(27)29 um. P/E=0.92. Vista polar circular de 19(26)29 um de didmetro. Exina de 1.3 um de grosor, con la nexina y sexina del mismo espesor, ornamentacién al ML microreticulada, al MEB con microespinas sobre los muros. Colpos delgados con terminaciones agudas. Poro circular de 2.6(3.2)3.9 um de largo x 2.6 um de ancho. Indice del drea polar 0.34, media. Rumex mexicanus Meisn. 1.5 Km al NE de San Andrés Metla, Estado de México. J. Elias 14] (ENCB). Lamina VII, Figuras 76-81. Polen tricolporado a tetracolporado, semitectado, esférico de 23(28)33 x 26(28)31 um. P/E=1. ‘Vista polar circular de 26(30)33 wm de didmetro. Exina de 1.3 wm de grosor, con la nexina de menor espesor que la sexina, ornamentaci6n al ML microreticulada, al MEB con microverrugas sobre los muros. Colpos delgados con Quiroz G. etal.: Pollen of Polygonaceae 15 terminaciones agudas. Poro lalongado de 3.9(4.9)6.5 um de largo x 2.3 um de ancho. Indice del d4rea polar 0.39, media. Rumex obtusifolius L. Alrededores de Villa del Carbén, Estado de México. J. Gonzdlez 892 (ENCB). Lamina VII, Figuras 82-87. Polen tricolporado, semitectado, oblato de 23(28)30 x 27(31)33 um. P/E=0.64. Vista polar circular de 27(30)35 wm de didmetro. Exina de 1.4 pm de grosor, con la nexina de menor espesor que la sexina, ornamentacién al ML y al MEB reticulada. Colpos delgados a veces en mayor numero (4 o 5), uniendose en uno de los polos (sincolpado). Poros lalongados de 2.6(3.7)3.9 um de largo x 3.8 wm de ancho. Indice del drea polar 0.23, pequefia. Rumex pulcher L. Terrenos del antigiio Lago de Texcoco, ca. Pefién de los Bafios, Estado de México. J. Rzedowski 2250 (ENCB). Lamina VIII, Figuras 88-96. Polen tncolporado, algunas veces tetracolporado, semitectado, suboblato de 26(29)33 x 28(33)37 um. P/E=0.87. Vista polar circular de 26(39)33 wm de didmetro. Exina de 1.6 um de grosor, con la nexina de menor espesor que la sexina, ornamentaci6n al ML y al MEB reticulada. Colpos delgados con terminaciones agudas. Poro lalongado de 3.9(4.3)5.2 um de largo x 3.9 um de ancho. Indice del area polar 0.21, pequefia. CLAVE PARA DIFERENCIAR LOS TAXA DELA FAMILIA POLY GONACEAE 1.- Polen tectado. 2.- Endoabertura de forma circular a lolongada. ................... Eriogonum jamesii 2.- Endoabertura lalongada a eliptica. 3.- Exina de mayor grosor en los polos. ................ Polygonum convolvulus 3.- Exina de igual grosor en todo el contorno.......... Polygonum argyrocoleon Polygonum aviculare 1.- Polen semitectado. 4.- Polen pen poradds.... «xsissivaivehaseeeel eens Polygonum hydropiperoides Polygonum lapathifolium Polygonum mexicanum Polygonum punctatiun 4.- Polen tricolporado, tetracol porado, sincolpado, pericolporado, o pericolpado. 5.- Polen microrreticulado, tricolporado, tetracolporado, algunas veces Sincolpado, i2xys.annamasags> dca doses. Ug vene OT ANGeh SRE Rumex acetocella Rumex conglomeratus Rumex crispus Rumex flexicaulis Rumex mexicanus 16 PHYTOLOGIA July 1998 volume 85(1):1-18 Rumex obtusifolia Rumex pulcher 5.- Polen ampliamente reticulado, pericolpado. ......... Polygonum amphibiurn Polygonum coccineum Polygonum hartwrightii DISCUSION Y CONCLUSIONES La variabilidad de los granos de polen de las Polygonaceae permite designar a esta © familia como euripalina, lo que en muchos casos ayuda a distinguir géneros e incluso especies por sus caracterfsticas polinicas. Asi, el polen de esta familia es un valioso auxiliar en estudios sistemdticos infragenéricos. Los granos de polen de los géneros Eriogonum, Polygonum, y Rumex se diferencian por el tipo de exina que presentan, tectada en Eriogonum y en P. argyrocoleon, P. aviculare, y P. convolvulus. Polen semitectado se observa en todo el género Rumex del Valle y en siete especies de Polygonum, estos géneros a su vez se diferencian por el tipo de ornamentacién y aberturas que tienen; asi Rumex es microrreticulado, 3 o 4 colporado a veces sincolpado, mientras que Polygonum es ampliamente reticulado y pericolpado o periporado. Las caracteristicas del polen de Eriogonum jamesii concuerdan con las que senala Wodehouse (1931), particularmente la exina es gruesa y bien estructurada, del tipo que se relaciona con la polinizaci6n entomédfila. E] género mejor representado en el Valle de México es Polygonum con diez especies cuyo polen muestra grandes variaciones en sus caracteristicas morfoldégicas como son: forma, tamafio, tipo, numero, y arreglo de aberturas, asi como la omamentaciOn. Al comparar los resultados obtenidos en este trabajo con los de van Leeuwen et al. (1988) se encontré que el tipo sefialado como Polygonum amphibium (reticulado, pantocolpado) se encuentra en la especie del mismo nombre presente en el Valle de México, ademas se observé en P. coccinum y P. hartwrightii. Las caracteristicas que permiten diferenciar a estas especies es el arreglo de los colpos en pentagonos regulares y la ornamentacidén constituida por un amplio reticulo. El polen de Polygonum aviculare y P. argyrocoleon coincide con el tipo P. aviculare (tectado, psilado, 4-zonocolporado, o pantocolporado) en lo referente a la gruesa nexina y la sexina con pequefias perforaciones sdlo visibles al MEB. Sin embargo, difiere en la presencia de microverrugas presentes en las especies aqui estudiadas. El polen de Polygonum convolvulus tiene las caracteristicas sefialadas para el tipo del mismo nombre (tectum sin perforaciones 0 con pequefias perforaciones, sexina engrosada en los polos, 3-zonocolporado) no relacionada con una _ polinizacién entomédfila. Quiroz G. etal.: Pollen of Polygonaceae 17 El tipo Polygonum persicaria (reticulado, pantoporado) se encontr6é en P. hydropiperoides, P. lapathifolium, P. mexicanum y P. punctatum. Wodehouse (1931) sugiere que las aberturas de los tipos Polygonum amphibium y P. persicaria, se arreglan de acuerdo a un sistema en el cual las aberturas estan situadas en la parte media de los lados de los pent4gonos que forman las caras de un dodecahedro pentagonal. Con este sistema, los granos de polen deben tener 30 aberturas. Sin embargo, en las especies estudiadas se observaron variaciones en el ntimero de aberturas, inclusive se aprecia una amplia variacion en el numero de colpos y poros, por lo que coincidimos con la opinién de Hedberg (1946) sobre un arreglo de aberturas mds bien desordenado. La variaci6n en el numero de aberturas del género Rumex es muy marcada, van Leeuwen et al. (1988) relaciona el numero de colpos con el nivel de poliploidia. den Nijs et af. (1980) con base en el numero cromosémico separa tres grupos: 1) Diploides, tienen una alta proporciédn de granos de polen 3-colporados y un numero reducido de pantocolpados. 2) Tetraploides varfan del 56 al 100% de polen 3-colporado. 3) Hexaploides tiene del uno al 99% de polen 3-colporado. Los resultados de los autores antes mencionados y los obtenidos en el presente estudio confirman que pocas especies de Rumex son exclusivamente 3-colporados, lo que indica diferentes grados de poliploidia. Las variaciones en el numero cromosémico del género Rumex y de Polygonum parecen estar relacionados con la forma de vida de estas plantas que corresponden a malezas perennes en donde se ha encontrado con frecuencia la poliploidia (Mulligan & Findlay 1970). En cuanto a endoaberturas se observa que las especies de Rumex que tienen polen principal mente 3-colporados, muestran 3-endoaperturas bien diferenciadas que pueden ser circulares, lalongadas o lolongadas, mientras en los taxa con mayor ntimero de colpos, se aprecia una reduccion en el numero de ellas, esta observacién concuerda con lo encontrado por van Leeuwen et al. (1988). Al relacionar la morfologia del polen con la clasificacién taxonémica de Melchior (1964) en las tres subfamilias que el autor reconoce, situa al género Eriogonum en la tribu Eriogoneae, de la subfamilia Eriogonoideae; a Polygonum en la Polygoneae y a Rumex en la tnbu Rumiceae de la Polygonoideae, mientras que la subfamilia Coccoloboideae carece de representantes en el Valle de México. Esta separacién coincide en gran parte con los tipos polinicos determinados. Sin embargo, es de hacer notar las variaciones tan marcadas que muestra el polen del género Polygonum en sus caracteristicas, raz6n por la cual autores como Hedberg (1946) propone una nueva clasificaci6n taxondémica basada en la morfologia del polen. 18 PHYTOLOGIA July 1998 volume 85(1):1-18 BIBLIOGRAFIA den Nijs, J.C.M., H. Hooghiemstra, & H. Schalk. 1980. Biosystematic studies of the Rumex acetosella complex (Polygonaceae). IV. Pollen morphology and the possibilities of identification of cytotypes in pollen analysis. Phyton 20:307-323. Erdtman, G. 1943. An Introduction to Pollen Analysis. Ronald Press, New York, New York. 239 pp. Erdtman, G. 1952. Pollen Morphology and Plant Taxonomy. Angiosperms. Almoavist & Wiksell, Stockholm, Sweden. 539 pp. Hedberg, O. 1946. Pollen morphology in the genus Polygonum L. s.lat. and its taxonomical significance. Sven. Bot. Tidskr. 40:371-404. Huang, T.C. 1972. Pollen Flora of Taiwan. National Taiwan University Botany Department Press, Taipei, Taiwan. 297 pp. Kapp, R.O. 1969. Pollen and Spores. W.M.C. Brown Company Publishers, Dubuque, lowa. 249 pp. Lawrence, M.G.H. 1969. Taxonomy of Vascular Plants. MacMillan Company, New York, New York. 823 pp. Markgraf, V. & H.L. D’Antoni. 1978. Pollen Flora of Argentina. University of Arizona Press, Tucson, Anizona. 208 pp. Melchior, H. 1964. A. Engler’s. Syllabus der Pflanzenfamilien 11. Gebriider Borntraeger, Berlin Nikolassee. 666 pp. Mulligan, G.A. & J.N. Findlay. 1970. Reproductive systems and colonization in Canadian weeds. Can. J. Bot. 48:859-860. Nowicke, J.W. & J.J. Skvarla. 1979. Pollen morphology: the potential influence in higher order systematics. Ann. Misouri Bot. Gard. 66:633-700. Palacios-Chavez, R., B. Ludlow Wiechers, & R. Villanueva. 1991. Flora palinoldégica de la reserva de la bidsfera de Sian Ka’an, Quintana Roo. Punt, W., S. Blackmore, S. Nilsson, & A. Le Thomas. 1994. Glossary of pollen and spore terminology. LPP. Contnbutions series No. 1. LPP Foundation, Utrecht, The Netherlands. 71 pp. Quiroz-Garcia, D., R. Palacios-Chdvez, & M.L. Arreguin-Sdnchez. 1995. Morfologfa de los granos de polen de las familias Polygonaceae y Sapindaceae de Chamela, Jalisco. Memorias del VIII Coloquio de Paleobotdnica y Palinologia, ENCB, IPN. México, D.F., México. Pp. 80-94. Roubik, D.W. & J.E. Moreno. 1991. Pollen and Spores of Barro Colorado Island. Missouri Botanical Garden, St. Louis, Missouri. 268 pp. Rzedowski, J. & G.C. de Rzedowski. 1979. Flora Fanerogamica del Valle de México. Compafia Editorial Continental, S.A., México, D.F., México. Volumen 1. 403 pp. van Leeuwen, P., W. Punt, & P.P. Hoen. 1988. Polygonaceae. Review of Palaeobotany and Palynology. 57:81-151. Wodehouse, R.P. 1931. Pollen grains in the identification and classification of plants VI. Polygonaceae. Amer. J. Bot. 18:749-765. Phytologia (July 1998) 85(1):19-21. NEW NAMES AND COMBINATIONS INASTERACEAE: HELIANTHEAE- ECLIPTINAE William A. Weber University of Colorado Museum, Campus Box 315, Boulder, Colorado 80309 U.S.A. ABSTRACT The following nomenclatural changes are provided for my contribution to the Flora North America Project. The new genera Agnorhiza and Scabrethia are segregated from Wyethia, and subspecific rank is proposed for races of Helianthella californica. Specific status is proposed for Balsamorhiza hookeri var. lanata. KEY WORDS: Agnorhiza, Balsamorhiza, Helianthella, Scabrethia, Asteraceae, Flora of North America AGNORHIZA Agnorhiza (Jepson) W.A. Weber, gen. nov. TYPE: Helianthus invenustus Greene, Pittonia 1:284. 1889. Balsamorhiza, section Agnorhiza Jepson, Fl. California 1077. 1925. Wyethia, section Agnorhiza (Jepson) W.A. Weber, Amer. Mid. Naturalist 35:416. 1946. The species of Agnorhiza differ from Wyethia in lacking basal leaves and in having clearly petiolate, cordate-ovate, usually strongly reticulate-veined cauline leaves, as opposed to the lanceolate ones of Wyethia, and long, slender caudices, stout and short in Wyethia. They are narrow endemics of the “Mother Lode” area of the western foothills of the Sierra Nevada of California. Agnorhiza bolanderi (A. Gray) W.A. Weber, comb. nov. BASIONYM: Balsamorhiza bolanderi A. Gray, Proc. Amer. Acad. Arts 7:356. 1868. Agnorhiza elata (H.M. Hall) W.A. Weber, comb. nov. BASIONYM: Wyethia elata H.M. Hall, Univ. California Publ. Bot. 4:208. 1912. 19 20 PHYTOLOGIA July 1998 volume 85(1):19-21 Agnorhiza invenusta (Greene) W.A. Weber, comb. nov. BASIONYM: Helianthus invenustus Greene, Pittonia 1:284. 1889. Agnorhiza ovata (Torrey & Gray) W.A. Weber, comb. nov. BASIONYM: Wyethia ovata Torrey & Gray ex Torrey in Emory, Mexican Boundary Survey 143. 1848. Agnorhiza reticulata (Greene) W.A. Weber, comb. nov. BASIONYM: Wyethia reticulata Greene, Bull. California Acad. Sci. 1:9. 1884. BALSAMORHIZA Balsamorhiza lanata (W.M. Sharp) W.A. Weber, comb. nov. BASIONYM: Balsamorhiza hookeri Nutt. var. lanata W.M. Sharp, Ann. Missoun Bot. Gard. 22:130. 1935. HELIANTHELLA Helianthella californica A. Gray subsp. nevadensis (Greene) W.A. Weber, comb. nov. BASIONYM: Helianthella nevadensis Greene, Bull. California Acad. Sci. 1(3):89. 1885. Helianthella californica A. Gray subsp. shastensis (W.A. Weber) W.A. Weber, comb. nov. BASIONYM: Helianthella californica A. Gray var. shastensis W.A. Weber, Amer. Midl. Naturalist 48:30. 1952. SCABRETHIA Scabrethia W.A. Weber, gen. nov. [for “scabrous Wyethia”]. TYPE: Wyethia scabra W.J. Hooker, London J. Bot. 6:245. 1847. Perennis, tota hispida et scabrida. Cauli numerosi, 1.5—4.0 dm alti; folia basalibus desunt; folia caulina linearibus, triplinervia. Capitulo solitario, 2 cm lato, phyllaria ad basin ovata, supra attenuata. Flores fertiles; flosculis radiorum 10-13, 3.5 cm longis. Cypselae glabrae; pappi coroniformi, arstae desunt. Stems numerous from underground caudices crowning a taproot, 1.54.0 dm high, very pale or white, hispid the entire length, as are all the vegetative parts, the long multicellular hairs becoming more dense toward the inflorescence, finally breaking off and leaving swollen bases which cause the surface of the plant to be very harshly scabrous. Leaves progressively reduced toward the base to about 2-3 cm Weber. New names in Ecliptinae 21 long, withering early; basal leaves absent. Leaves largest at the middle of the stem, linear, very stiff and harsh, entire, sessile, 3-15 cm long, 3-17 mm wide, abruptly narrowed or truncate at the base, mucronate at the apex, the nerves very pale, the lateral ones confluent toward the margin, giving the leaves a tniple-nerved appearance. Heads solitary, terminal, ca. 2 cm wide. Involucre hemispherical, the outer series of phyllaries narrowly lanceolate to linear, attenuate from broad, ovate bases, pubescent mostly on the margins; inner series broader, rigid, ovate-lanceolate with attenuate tips, the trichomes finer, more appressed, and covering the entire surface. Disk-flowers glabrous. Ray-flowers 10-13, ca. 3.5 cm long, lemon-yellow, rather densely pubescent on the tube and dorsally along the nerves. Paleae deeply carinate above, pubescent on the midribs and margins. Achenes glabrous, 6-8 mm long. Pappi low, coroniform, lacking conspicuous awns. One complete plant of subspecies scabra was excavated at the Moffat County, Colorado site. The main root body was 12 cm long, 3 cm wide, with loose, flaking bark. A few slender, elongate lateral branches 0.5 cm wide, anise from this. The main root forks below, producing two branches averaging 2.5 cm wide and 6 dm long. Distally, the root gives rise to two caudices 2 cm wide and 5 cm long; these in tum produce 2-4 smaller caudices which give rise just below ground level into 4-6 still smaller ones. From these caudices anise 30-40 aerial stems. The scabrous pubescence consists of 3-celled trichomes curved toward the leaf and stem apex; the terminal cell is thick-walled, smooth, rigid, and sharply pointed. The two basal cells are isodiametric, roughly spherical, and papillose. Most of the trichomes lack the terminal cell, which results in the extreme scabrosity. The basal cell of each trichome is surrounded by a rosette of bulliform cells distinctly larger than the adjacent epidermal cells. Stomata are extremely numerous. Scabrethia scabra (W.J. Hooker) W.A. Weber, comb. nov. BASIONYM: Wyethia scabra W.J. Hooker, London J. Bot. 6:245. 1847. Scabrethia scabra (W.J. Hooker) W.A. Weber subsp. attenuata (W.A. Weber) W.A. Weber, comb. nov. BASIONYM: Wyethia scabra W.J. Hooker var. attenuata W.A. Weber, Amer. Midl. Naturalist 35:425. 1946. Scabrethia scabra (W.J. Hooker) W.A. Weber subsp. cinerea (W.A. Weber) W.A. Weber, comb. nov. BASIONYM: Wyethia scabra W.J. Hooker var. cinerea W.A. Weber, Amer. Midl. Naturalist 35:425. 1946. Phytologia (July 1998) 85(1):22-31. WETLAND STATUS OF HILLSIDE BOGS, WETLAND PINE SAVANNAS, AND MUCK BOGS IN THE WEST GULF COASTAL PLAIN Michael H. MacRoberts & Barbara R. MacRoberts Bog Research, 740 Columbia, Shreveport, Louisiana 71104 U.S.A. & Herbarium, Museum of Life Sciences, Louisiana State University in Shreveport, Shreveport, Louisiana 71115 U.S.A. ABSTRACT The wetland status of West Gulf Coastal Plain hillside bogs, wetland pine savannas, and muck bogs is apparently equivocal. We examined regulatory and nonregulatory wetland definitions and found that these plant communities conform to all wetland criteriaand therefore should be recognized as wetlands. KEY WORDS: Hillside bog, wetland pine savanna, muck bog, wetlands, pitcher plants INTRODUCTION In pre-European times, bogs and wetland pine savannas covered two to four million acres of the West Gulf Coastal Plain (WGCP). These were open areas with generally low tree densities, high water tables, and a rich predominately heliophilous, hydrophytic herbaceous ground cover (Bridges & Orzell 1989; Harcombe et al. 1993). Today, only one to five percent of the onginal acreage remains, most of which is badly degraded. These communities once covered extensive acreage in the East Gulf Coastal Plain and the Atlantic Coastal Plain as well (Brewer 1998; Folkerts 1982, 1991; Hermann 1995). Having recently completed floristic inventories on bogs and wetland pine savannas in the WGCP (MacRoberts & MacRoberts 1988, 1990, 1991, 1992, 1993, 1998a, 1998b), we were surprised to find that the wetland status of these communities is uncertain, even though there are clear jurisdictional/regulatory and scientific definitions that point to their being wetlands. The U.S.. Army Corps of Engineers and the U.S. Fish and Wildlife Service provide regulatory definitions and procedures for identifying wetlands (Cowardin ef 22 oe MacRoberts & MacRoberts: West Gulf Coastal Plain wetlands 23 al. 1979; U.S. Army Corps of Engineers Delineation Manual 1987 [COE Manual 1987]; Federal Manual for Identifying and Delineating Jurisdictional Wetlands 1989 (Federal Manual 1989]; Fretwell et al. 1996; National Research Council 1995 [NRC 1995]; Policansky 1998; Reed 1988 and 1996 update; U.S. Soil Conservation Service 1991 [U.S. Soil 1991]; Tiner 1988, 1996). Three criteria are used to designate wetlands: hydrology, soils, and vegetation. The National Wetland Inventory (1974- 1987) has produced maps showing where wetlands occur. In this paper we address the wetland status of WGCP hillside bogs, wetland pine savannas, and muck bogs. Detailed descriptions of these and similar sites can be found elsewhere (Bridges & Orzell 1989; Harcombe et al. 1993; Kral 1955; Lodwick 1975; MacRoberts & MacRoberts 1988, 1990, 1991, 1992, 1993, 1998a, 1998b; Nixon & Ward 1986; Rowell 1949; Streng & Harcombe 1982). We will be concerned especially with plants since we have conducted floristic inventones of bogs and wetland pine savannas. STUDY SITES Our sample consists of seventeen bogs and wetland pine savannas in the WGCP: thirteen hillside bogs, two wetland pine savannas, and two muck bogs. The locations of the seventeen sites are shown in Figure 1. Andrew’s and Chester’s muck bogs (A- MB, C-MB) are on the Gus Engeling Wildlife Management Area, Anderson County, Texas. Cooter’s hillside bog (C-HB) is on the Vernon Ranger District of the Kisatchie National Forest, Vernon Parish, Louisiana. Fixit, Middle Branch, Vine, Penrod, RCW, Robin, Sparrow, Woodcock, 360A, and 360B hillside bogs (NATCH) are on the Kisatchie Ranger District of the Kisatchie National Forest, Natchitoches Parish, Louisiana. Strange Road hillside bog (SR-HB) is on the Winn Ranger District of the Kisatchie National Forest, Natchitoches Parish, Louisiana. Turkey Creek and Lance Rosier wetland pine savannas (TC-WPS, LR-WPS) occur on the Big Thicket National Preserve, Hardin and Tyler counties, Texas. “TX-HB” hillside bog is a composite created by combining various incomplete lists taken from Nixon & Ward (1986) and Orzell (1990) and our personal observations on a group of hillside bogs centering on Boykin Springs, Angelina National Forest, Angelinaand Jasper counties, Texas. The composite is an attempt to produce a representative hillside bog floristics for this region of east Texas, comparable to our other sites. Except for TX-HB, we conducted year-round floristic inventories on each bog or wetland pine savanna in the sample. Each was visited every two to four weeks from March through November. We developed complete plant lists for these sites. That is, our surveys were designed to identify all the vascular plant species that grew within each bog or wetland pine savanna. Individual site plant lists range from 64 species at the very small Robin hillside bog (0.03 ha) to 131 species at the large Cooter’s hillside bog (3.0 ha). The mean site, which measured between 0.4 and 0.8 ha, had 98 species. Soil maps are available for all sites (Coffee 1975; Deshotels 1987; Martin 1990), but many sites are so small as not to have been mapped separately from the surrounding landscape. We have made observations on hydric conditions at all sites. MacRoberts & MacRoberts: West Gulf Coastal Plain wetlands 25 DEFINITIONS The Federal Manual (1989) defines wetlands as having “. . . one or more of the following three attributes: (1) at least periodically, the land supports predominately hydrophytes, (2) the substrate is predominately undrained hydric soil, and (3) the substrate is nonsoil and is saturated with water or covered by shallow water at some time during the growing season of each year” (see also Cowardin et al. 1979). U.S. Soil (1991) and the COE Manual (1987) define wetlands very similarly (Tiner 1996; NRC 1995). However, the COE Manual (1987) states that all three criteria, not just “one or more,” must be met before a site can be delineated as a wetland. While the NRC (1995) provides a “scientific” -- not a “regulatory” -- definition of wetlands (Policansky 1998), it stipulates virtually the same criteria and repeats the tno: recurrent, sustained inundation or saturation at or near the surface; hydric soils; and hydrophytic vegetation, but emphasizes that hydrophytic vegetation, by definition, indicates hydric soils. Below, we examine each of the criteria used in these definitions to determine whether or not WGCP bogs and wetland pine savannas are classifiable as wetlands. A history of wetlands in the United States is given by Vileisis (1997) and a history of the involvement of the Corps of Engineers in water matters in the United States is given in Shallat (1994). VEGETATION Vegetation appears to be the defining criterion for wetlands that is most emphasized by all agencies. The U.S. Fish and Wildlife Service has produced a national plant list that ranks species on a wetland (hydrophytic) gradient (Reed 1988, 1996 update). According to the Federal Manual (1989:5): An area has hydrophytic vegetation when, under normal circumstances: (1) more than 50 percent of the composition of the dominant species from all strata are obligate wetland (OBL), facultative wetland (FACW), and/or facultative (FAC) species, or (2) a frequency analysis of all species within the community yields a prevalence index value of less than 3.0 (where OBL = 1.0, FACW = 2.0, FAC = 3.0, FACU = 4.0, and UPL = 5.0). Reed (1988) defines these five categories of plant species: 1) obligate wetlands (OBL): greater than 99% occurrence in wetlands; 2) facultative wetland (FACW): 67% - 99% occurrence in wetlands, but occasionally found in nonwetlands; 3) facultative (FAC): equally likely to occur in wetlands or nonwetlands (estimated probability 34% - 66%); 4) facultative upland (FACU): usually occur in nonwetlands (estimated probability 67% - 99%), but occasionally found in wetlands; and 5) obligate uplands (UPL): occurs almost always (estimated > 99%) in nonwetlands. 26 PHYTOLOGIA July 1998 volume 8X 1):22-31 Table 1 compares percentage frequency of all plants from our sample of seventeen sites as rated by Reed’s (1988) hydrophytic classification. Table 1 also gives the prevalence index and the total number of species at each site. We compared the 1996 update of Reed’s listing with the 1988 original and found the two to be virtually identical. Since there is no significant difference between the two lists, we use Reed (1988). Table 1. Species percents by wetland gradient categories, prevalence index, and sample size (HB = hillside bog, MB = muck bog, WPS = wetland pine savanna). OBL | FACW] FAC] FACU [ UPL [| Prev. | Sample 0 eee [Andrew’sMB_ | 6O {| 31 | 9 | 0 | Chester'sMB- | 58 | 30 | 10 | 1 | 1 | 15) ) aa E40. | |} 710. | 1) oak Rope ps Pe , Penrod HB | 36[ 38 | 17 |_7_] 2 | 20) | Sparow HS | 38.[ 31] 2] 11 | 0 | 203 | oom [Woodcock HB| 36] 37 | 18] 8 | 1 | 205 | 99 | 00: a eS A SO AO The flora of these sites is clearly dominated by hydrophytes. Between 88% and 100% of species at each site are OBL, FACW, and FAC (50% or higher qualifies a site as wetland). The prevalence index ranges from 1.49 to 2.17 (3.0 and lower qualifies as wetlands). The few FACU or UPL species that occur in these sites are either intruders taking advantage of microhabitat, are species misclassified by Reed (1988, 1996 update), or are species with wide hydric tolerance. For example, in the WGCP, Marshalliatenuifolia Raf. probably should be classified as a FACW species, and Pinus palustris Mill., Pityopsis graminifolia (Michx.) Nutt., and Liatris pycnostachya Michx. have a wide hydric tolerance that ranges from saturated to xeric soils (Bridges & Orzell 1989; MacRoberts & MacRoberts 1993, 1996; Marks & Harcombe 1981). MacRoberts & MacRoberts: West Gulf Coastal Plain wetlands ay No matter: A few disputable assignments make no difference. The figures are clear. All these sites are unquestionably vegetationally wetlands. Finally, if any additional evidence of the hydrophytic nature of the vegetation of hillside bogs, wetland pine savannas, and muck bogs be needed, the only sources required to identify virtually all vascular plants occurring in WGCP bogs and wetland pine savannas are either Godfrey & Wooton’s (1979, 1981) Aquatic and Wetland Plants of the Southeastern United States or Correll & Correll’s (1972) Aquatic and Wetland Plants of Southwestern United States. WATER The main hydrological criterion for wetlands is “permanent or periodic inundation, or soil saturation, for a significant period (usually a week or more) during the growing season” (Federal Manual 1989:15). Wetland pine savannas are poorly to very poorly drained and have high water tables. From winter through spring, they have standing, shallow water most of the time and penodically throughout the growing season (Bridges & Orzell 1989; Deshotels 1978; Harcombe et al. 1993; MacRoberts & MacRoberts pers. obs.; Streng & Harcombe 1982) although there are droughty periods in the summer and early autumn when the water table lowers. Both hillside bogs and muck bogs are saturated year round: the former from lateral movement of water from higher ground; the latter from both lateral movement of water, but also from standing water (MacRoberts & MacRoberts pers obs.). SOILS U.S. Soil (1991:1) defines a hydric soil as “. . . a soil that is saturated, flooded, or ponded long enough during the growing season to develop anaerobic conditions in the upper part” (see also Federal Manual 1989:6; Tiner 1996). U.S. Soil (1991) and the COE Manual (1987) list soils in the United States by series and subgroup considered to be hydric. The soils upon which hillside bogs, wetland pine savannas, and muck bogs occur -- Guyton, Jasco, Waller, Plank, Bienville, Anacoco, Wehadkee, and Nahatche (vertic Albaqualfs, typic Glossaqualfs, typic Fragiaqualfs, psammentic Paleudalfs, aeric and typic Fluvaquents) -- are listed as hydric (U.S. Soil 1991; COE Manual 1987). NATIONAL WETLANDS INVENTORY MAPS As part of this investigation we examined the most recent National Wetlands Inventory Maps (1974-1987) to see how the sites in our sample of hillside bogs, wetland pine savannas, and muck bogs were delineated. Except for the two muck bogs, none of the sites was designated as wetland. What the maps delineate as 28 PHYTOLOGIA July 1998 volume 85(1):22-31 wetlands are ponds, major streams, and those locations on topographic maps marked by “swamp” symbols; the remaining land is marked “upland.” Thus, if an area was not obviously a wetland by the fact of open water or was clearly marked as swamp on a topographic map, it was designated an upland. DISCUSSION By all agency criteria, hillside bogs, wetland pine savannas, and muck bogs are wetlands. Using the Cowardin et al. (1979) and Tiner (1988) “keys” for determining wetlands, WGCP wetland pine savannas and hillside bogs are palustrine, persistent, emergent, saturated wetlands; muck bogs are palustrine scrub-shrub or emergent wetlands. One of the peculiarities of the Corps of Engineers methodology that is often lost sight of is the three factor requirement whereby criteria established for vegetation, soils, and water must all be met (COE Manual 1989). However, as NRC (1995) points out, what this requirement overlooks is the strong causal relationship among water, soils, and vegetation. For example, plant communities dominated by OBL and FACW species develop only where the prevailing hydrologic regime is one in which flooding and saturation is frequent or extended. In fact, the situation is even more elementary since water, soils, and vegetation are interdependently defined. Hydrophytic vegetation, by definition, is associated with water; it is called hydrophytic because of its aquatic or wetland association. Thus, a strict requirement that soils, hydrology, and vegetation all meet “wetland” criteria is redundant at best, obfuscatory at worst, and becomes an exercise in excess or redundancy (see NRC 1995: 139-140.). Most definitions of wetlands establish sampling procedures that prove to be ambiguous (e.g., COE Manual procedures). We solved the problems arising from ambiguously defined “dominant species” or “stratum” by surveying year round and developing a total floristic list. In this way, all species that grow in a specific site are included and bias resulting from seasonal sampling or sampling that favors large, conspicuous, showy, or easily identified species such as longleaf pine or pitcher plants is avoided. Rare, small species, such as Bartonia verna (Michx.) Muhl. and Burmannia capitata (Gmel.) Mart., and species difficult to identify, such as members of the genera Rhynchospora and Carex, that are excellent indicators of hydrological conditions are not discriminated against in favor of species that are easily identified but which are not particularly good hydrological indicators. While the agencies have produced universal lists of hydric soils and hydrophytic plants, they have not produced universal keys to or lists of wetland plant communities. This means that at each site, individual plants must be collected and analyzed instead of using a simple key to decide community. Such keys have been developed locally by Marks & Harcombe (1981) and Cowardin et al. (1979), which can be employed with almost no detailed knowledge of the vegetation. Why there is ambiguity in classifying bogs and wetland pine savannas is difficult to understand. We conclude that itis due to the three-factor requirement of the Corps of Engineers, coupled with the fact that few individuals are botanically sophisticated | MacRoberts & MacRoberts: West Gulf Coastal Plain wetlands 29 enough to assess the local flora, either by sampling it as prescribed in the manuals or by conducting a year-round inventory involving the identification of all species. It is interesting that wetland pine savannas in the Carolinas, Georgia, and Florida are recognized as wetlands, yet these same communities in Alabama, Mississippi, Louisiana, and Texas are not (Fretwell et al. 1996). Suffice it to say that far larger areas of our landscape are wetlands than are so designated on wetland inventory maps. ACKNOWLEDGMENTS Financial support was provided in part by a USGS Biological Resources Divisions Species at Risk grant to the National Wetlands Research Center. Carla Clark and Gene Romine, Noel Memorial Library, Louisiana State University in Shreveport, helped in obtaining numerous documents. Joe Liggio aided with soils and orchids. Steve Orzell, Avon Park Air Force Range, Florida, and Susan L. Grace, National Wetlands Research Center, Lafayette, Louisiana, read an earlier version of this paper. LITERATURE Brewer, J.S. 1998. Patterns of plant species richness in a wet slash pine (Pinus elliottii) savanna. J. Torrey Bot. Soc. 125:216-224. Bridges, E.L. & S.L. Orzell. 1989. Longleaf pine communities of the west gulf coastal plain. Natural Areas Journal 9:246-263. Coffee, D.R. 1975. Soil Survey of Anderson County, Texas. U.S. Soil Conservation Service, Washington, D.C. Correll, D.S. & H.B. Correll. 1972. Aquatic and Wetland Plants of the Southwestern United States. Stanford University Press, Palo Alto, California. Cowardin, L.M., V. Carter, F.C. Golet, & E.T. LaRoe. 1979. Classification of wetlands and deepwater habitats of the United States. U.S. Fish and Wildlife Service Report FWS/OBS-79/31. Washington, D.C. Deshotels, J.D. 1978. Soil Survey for Big Thicket National Preserve: Texas. United States Department of Agriculture, Washington, D.C. Federal Interagency Committee for Wetland Delineation [Federal Manual]. 1989. Federal Manual for Identifying and Delineating Jurisdictional Wetlands. U.S. Army Corps of Engineers, U.S. Environmental Protection Agency, U.S. Fish and Wildlife Service, and U.S. Soil Conservation Service, Washington, D.C. Folkerts, G.W. 1982. The gulf coast pitcher plant bogs. American Scientist 70:260- 207. Folkerts, G.W. 1991. A preliminary classification of pitcher plant habitats in the ’ southeastern United States. J. Alabama Acad. Sci. 62: 199-225. Fretwell, J.D., J.S. Williams, & P.J. Redman (compilers). 1996. National Water Summary on Wetland Resources. U.S. Geological Survey, Water-Supply Paper 2425, Washington, D.C. 30 PHYTOLOGIA July 1998 volume 85( 1):22-31 Godfrey, R.K. &J.W. Wooten. 1979. Aquatic and Wetland Plants of Southeastern United States. Monocotyledons. University of Georgia Press, Athens, Georgia. Godfrey, R.K. &J.W. Wooten. 1981. Aquatic and Wetland Plants of Southeastern United States. Dicotyledons. University of Georgia Press, Athens, Georgia. Harcombe, P.A., J.S. Glitzenstein, R.G. Knox, S.L. Orzell, & E.L. Bridges. 1993. Vegetation of the longleaf pine region of the west gulf coastal plain. Proc. Tall Timbers Fire Ecology Conf. 18:83-104. Hermann, S.M. 1995. Status and management of Florida’s camivorous plant communities. Florida Game and Fresh Water Fish Commission Nongame Wildlife Program Project Report. Tallahassee, Flonda. Kral, R. 1955. A floristic comparison of two hillside bog localities in northeastern Texas. Field and Lab. 23:47-69. Lodwick, L.N. 1975. Net aerial primary production of three east Texas peat bogs. MS thesis: Baylor University, Waco, Texas. MacRoberts, B.R. & M.H. MacRoberts. 1988. Floristic composition of two west Louisiana pitcher plant bogs. Phytologia 65:184-190. MacRoberts, B.R. & M.H. MacRoberts. 1990. Vascular flora of two west Louisiana pitcher plant bogs. Phytologia 68:271-275. MacRoberts, B.R. & M.H. MacRoberts. 1991. Floristics of three bogs in western Louisiana. Phytologia 70: 135-141. MacRoberts, B.R. & M.H. MacRoberts. 1992. Floristics of four small bogs in western Louisiana with observations on species-area relationships. Phytologia 73:49-S6. MacRoberts, B.R. & M.H. MacRoberts. 1993. Floristics of a bog in Vernon Parish, Louisiana, with comments on noteworthy bog plants in westem Louisiana. Phytologia 75:247-258. MacRoberts, B.R. & M.H. MacRoberts. 1996. Floristics of xeric sandhills in east Texas. Phytologia 80:1-7. MacRoberts, B.R. & M.H. MacRoberts. 1998a. Floristics of wetland pine savannas in the Big Thicket National Preserve, southeast Texas. Phytologia 85(1):40-50. MacRoberts, B.R. & M.H. MacRoberts. 1998b. Floristics of muck bogs in east central Texas. Phytologia 85(1):61-72. Marks, P.L. & P.A. Harcombe. 1981. Forest vegetation of the Big Thicket, Southeast Texas. Ecol. Monog. 51:287-305. Martin, P.G. 1990. Soil Survey of Natchitoches Parish, Louisiana. U.S. Soil Conservation Service, Washington, D.C. National Research Council [NRC]. 1995. Wetlands: Characteristicsand Boundaries. National Academy Press, Washington, D.C. National Wetland Inventory. 1974-87. U.S. Dept. of Interior. U.S. Fish and Wildlife Service. Maps. Nixon, E.S. & J.R. Ward. 1986. Floristic composition and management of east Texas pitcher plant bogs. Pp. 283-287. In: D.L. Kulhavy & R.W. Conner (eds.), Wilderness and Natural Areas in the Eastern United States: A Management Challenge. Center for Applied Studies, School of Forestry, Stephen F. Austin State University, Nacogdoches, Texas. Orzell, S.L. 1990. Texas Natural Heritage Program Inventory of National Forests and Grasslands in Texas. Texas Parks and Wildlife Department, Austin, Texas. Policansky, D. 1998. Science and decision making for water resources. Ecol. Applications 8:610-618. MacRoberts & MacRoberts: West Gulf Coastal Plain wetlands 31 Reed, P.B. 1988 (and 1996 update). National List of plant species that occur in wetlands: national summary. U.S. Fish and Wildlife Service, Biological Report 88(24). Washington, D.C. Rowell, C.M. 1949. Floral composition of a sphagnum bog in Robertson County, Texas. MS thesis, Texas A&M University, College Station, Texas. Shallat, T. 1994. Structures in the Stream: Water, Science, and the Rise of the U.S. Army Corps of Engineers. University of Texas Press, Austin, Texas. Streng, D.R. & P.A. Harcombe. 1982. Why don’t east Texas savannas grow up into forests? Amer. Midl. Naturalist 108:278-294. Tiner, R.W. 1988. Field Guide to Nontidal Wetland Identification. Maryland Department of Natural Resources, Annapolis Maryland and U.S. Fish and Wildlife Service, Newton Corer, Maryland. Tiner, R.W. 1996. Wetland definitions and classifications in the United States. Pp. 27-34. In: J.D. Fretwell, J.S. Williams, & P.J. Redman (compilers). 1996. National Water Summary on Wetland Resources. U.S. Geological Survey, Water-Supply Paper 2425, Washington, D.C. U.S. Army Corps of Engineers [COE Manual]. 1987. Corps of Engineers wetlands delineation manual. Tech. Report Y-87-1. U.S. Army Corps of Engineers, Waterways Experiment Station, Vicksburg, Mississippi. U.S. Soil Conservation Service [U.S. Soil]. 1991. Hydric Soils of the United States. U.S. Soil Conservation Service, Washington, D.C. Vileisis, A. 1997. Discovering the Unknown Landscape: A History of America’s Wetlands. Island Press, Washington, D.C. Phytologia (July 1998) 85(1):32-39. COMMUNITY CLASSIFICATION OF WEST GULF COASTAL PLAIN BOG COMMUNITIES: A FLORISTIC ASSESSMENT iichsct Tr Mackobers @ Barta R MacRabets Bog Research, 740 Columbia, Shreveport, Louisiana 71104 U.S.A. & Museum of Life Sciences, Louisiana State University in Shreveport, One University Place, Shreveport, Louisiana 71115 U.S.A. ABSTRACT Many community classifications separate West Gulf Coastal Plain hillside bogs, wetland pine savannas, and muck bogs. Based on flonstic data, hillside bogs and wetland pine savannas are similar and muck bogs are different. KEY WORDS: Hillside seepage bog, wetland pine savanna, pitcher plant bog, muck bog, West Gulf Coastal Plain INTRODUCTION In the course of our work on bogs in the West Gulf Coastal Plain (WGCP), we repeatedly have encountered the problem of classifying bog communities (Bridges & Orzell 1989; Diamond et al. 1987; Folkerts 1991; Louisiana Natural Hentage Program 1988; Peet & Allard 1993; Texas Natural Hentage Program 1993; Texas Organization of Endangered Species 1992; Weakley et al. 1998). Received opinion is that at least two communities are represented (Bridges & Orzell 1989; Diamond et al. 1987; Louisiana Natural Heritage Program 1988), but in the East Gulf Coastal Plain (EGCP), bogs (or their counterparts) are considered to be the same plant community or at least members of a plant community type (Folkerts 1991). However, Harcombe et al. (1993) cautioned that in the WGCP these assessments have preceded a thorough investigation, certainly a quantitative one. Our studies of the floristics of hillside bogs (MacRoberts & MacRoberts 1988, 1990, 1991, 1992, 1993) and our cursory observations of wetland pine savannas, and savanna and muck bog descriptions in the literature (Bridges & Orzel] 1989; Harcombe et al. 1993; Kral 1955; Lodwick 1975; Rowell 1949), have led us to reexamine the traditional bog/savanna classifications. The purpose of our research was not to 32 MacRoberts & MacRoberts: West Gulf Coastal Plain bog communities 33 adjudicate on community classification, but to obtain floristic information about plant communities. To this end, we made year-round floristic surveys of muck bogs and wetland pine savannas (MacRoberts & MacRoberts 1998a, 1998b). Floristic lists from each site are directly comparable and therefore provide comparative information about communities. METHODS In the course of the past twelve years, we have developed total floristic lists for twelve hillside bogs on the Kisatchie National Forest, Natchitoches and Veron parishes, Louisiana; two wetland pine savannas on the Big Thicket National Preserve, Hardin and Tyler counties, Texas; and two muck bogs on the Gus Engling Wildlife Management Area, Anderson County, Texas (MacRoberts & MacRoberts 1988, 1990, 1991, 1992, 1993, 1998a, 1998b). Using the lists of Bndges & Orzell (1989), Orzell (1990), Nixon & Ward (1986), and our own observations since 1993, we also have developed a plant list for Texas hillside bogs. This list centers on the Angelina National Forest in Angelina and Jasper counties. Although the list is not based on a year-long survey of a single site, it is probably fairly representative of a typical Texas hillside bog. In addition to these data, there are variously complete floristic lists that allow some comparisons among bog types (Ajilvsgi 1979; Allen et al. 1988; Bridges & Orzell 1989; Kral 1955; Lodwick 1975; Nixon & Ward 1986; Orzell 1990; Rowell 1949; Streng & Harcombe 1982; Texas Natural Heritage Program 1993; Watson 1979, 1982). There are many objective methods for classifying communities: multifactor ecological classification systems use both biotic and abiotic factors (Turner & Van Kley in prep.); others emphasize floristic data (Weakley et al. 1998). We have used a floristic approach, which harkens back to the European phytosociological tradition in which plant communities were recognized by their vegetation as a whole, of which some plants were deemed more sensitive expressions of a given relationship than others (Fauth et al. 1996; van der Maarel 1975; McIntosh 1975; Whittaker 1975; Wilson et al. 1996). Following the lead of Nixon & Ward (1986), we have used Sorensen’s Index of Similarity, one of the older methods of community comparison (Sorensen 1948). The formula is IS = (2C/A+B) x 100, where C is the number of species common to the two samples, A is the total number of species in sample A, and B is the total number of species in sample B. Interpretation is relatively simple: 100 means that the compared communities (or samples) have identical taxa, and 0 means that they have no taxain common. Figures in between require some interpretation: the higher the number, the greater the similarity. We have found that the IS of samples of the same community ranges from about 55 to 60 and up. The criticism that the Index overvalues rare species is unjust (Southwood 1966); rather, the Index levels the field by not overemphasizing so-called “dominant” (large) generalists or otherwise conspicuous species. Further, the question addressed by the statistic is not whether or not there are different communities, but rather how similar the two samples are. 34 PHYTOLOGIA July 1998 volume 85( 1):32-39 Figure 1. Distribution of study Sites. See Table 1 for further locality information. MacRoberts & MacRoberts: West Gulf Coastal Plain bog communities 35 RESULTS Using Sorensen’s Index of Similarity, Table 1 compares the floristic similarity among two wetland pine savannas (LR-WPS and TC-WPS) (MacRoberts & MacRoberts 1998a), two muck bogs (A-MB and C-MB) (MacRoberts & MacRoberts 1998b), and two hillside bogs (LA-HB and TX-HB) (MacRoberts & MacRoberts 1993, unpublished observations; Orzell 1990; Nixon & Ward 1986). The six samples consist of between 105 and 131 species. Figure 1 shows the distribution of these sites. Table 1. Comparison of bogs and savannas by Sorensen’s Index of Similarity (IS). A-MB = Andrew’s Bog, Gus Engeling Wildlife Management Area in Anderson County, Texas; C-MB = Chester’s Bog, Gus Engeling Wildlife Management Area in Anderson County, Texas; LR-WPS = Lance Rosier Wetland Pine Savanna, Big Thicket National Preserve in Hardin County, Texas; TC-WPS = Turkey Creek Wetland Pine Savanna, Big Thicket National Preserve in Tyler County, Texas; LA-HB = Cooter’s Hillside Bog, Kisatchie National Forest in Vernon Parish, Louisiana; TX- HB = Texas Hillside Bog, Angelina National Forest in Angelina and Jasper counties, Texas. LR-WPS TC-WPS LA-HB PA BOS TERI REE rt tS! eee ee SAO GY CE ENS Pe SSS See EN eee ea ee eA a a DISCUSSION In this paper we have emphasized floristic data in the classification of bog communities. We do so not only because communities are about plants, but also because we had already collected complete floristic lists for hillside bogs. Comparisons of these plant lists show that hillside bogs and wetland pine savannas are basically the same floristically: at least in terms of total species. Muck bogs are distinct. There is a literature with partial floristic lists for wetland pine savannas and hillside bogs. For example, the Texas Natural Heritage Program (1993) describes hillside bogs and wetland pine savannas as two communities (“Longleaf Pine-Beakrush Series” and “Sphagnum-Beakrush Series”) and presents a short but essentially 36 PHYTOLOGIA July 1998 volume 85( 1):32-39 identical plant list for each. The Louisiana Natural Heritage Program (1988) calls these two communities “Hillside Bog” and “Pine Savanna,” but the lists of plants for the two is almost identical. Bridges & Orzell (1989) list the percent frequency of prevalent species for hillside bogs and for wet savannas. Of the 47 species listed in their wet savannas, 44 (94%) occur in hillside bogs (MacRoberts & MacRoberts 1988, 1990, 1991, 1992, 1993; Nixon & Ward 1986). Streng & Harcombe (1982) present a partial list of species for a wetland pine savanna in southern Tyler County that is basically identical with lists from hillside bogs farther north. Bridges & Orzell (1989) make the point that species prevalence is different between hillside bogs and wetland pine savannas. We can say little about prevalence since we have studied only a few sites. But on the basis of unquantified observations, Bridges & Orzell are probably correct; variation in abundance is something that should be studied in the bog community complex. We have emphasized floristics in these comparisons. Ecological classifications are beginning to be developed for WGCP plant communities, but these as yet do not include samples of all bog types (Harcombe et al. 1993; Turner & Van Kley in prep.; Van Kley in press). When more research has been completed, it will be interesting to see the similarities and differences among classifications that are based on different criteria. Why hillside bogs and wetland pine savannas have been considered separate plant communities in the WGCP is not entirely clear, but there may be several reasons. One may be that the floristics of most WGCP plant communities is poorly known and quantitative data are usually unavailable. Consequently, comparisons between and among communities have remained largely subjective. Another factor is that single species are often used as community indicators. In the case of hillside bogs, the presence or absence of Sarracenia alata Wood. appears to be important in classifying sites. In the WGCP, Sarracenia drops out of the flora 80 to 100 km inland, south of which much of the wetland pine savanna occurs. However, many hillside bogs also lack Sarracenia (Hermann 1990, 1995; Kral 1955; MacRoberts & MacRoberts 1991); for example, only 22% of hillside bogs in Natchitoches Parish have pitcher plants; whereas over 90% of those in Vernon Parish do (MacRoberts & MacRoberts pers. obs). Yet, hillside bogs without pitcher plants are not otherwise floristically different from sites with pitcher plants (MacRoberts & MacRoberts 1991). In the post oak region of Texas, Kral (1955) studied bogs without pitcher plants; whereas those studied by Rowell (1949) and Lodwick (1975) had them; all these sites were floristically similar. Consequently, while it is interesting that Sarracenia drops out of the WGCP flora at the Beauregard/Calcasieu Parish line in Louisiana and at the Tyler/Hardin County line in Texas, the floristic complement typical of bogs remains. Apparently because pitcher plants extend all the way to the coast in the EGCP (Folkerts 1991), a hillside bog/wetland pine savanna distinction is not made for bog communities in that region. As Folkerts (1982:260) said of pitcher plant habitat, “bogs in areas of little relief are typically called savannas.” The following scheme approximates our floristic findings for bog communities in the WGCP: WGCP Bog Communities A. Bog -savanna MacRoberts & MacRoberts: West Gulf Coastal Plain bog communities 37 1. Hillside bog 2. Wetland pine savanna B. Muck bog ACKNOWLEDGMENTS Persons and supporting institutions aiding with this research are acknowledged in our previous publications. Partial support was provided also by a USGS Biological Resources Division Species at Risk Grant to the National Wetlands Research Center, Lafayette, Louisiana. Steve Orzell, Avon Park Air Force Range, Florida, and Susan L. Grace, National Wetlands Research Center, Lafayette, Louisiana, made helpful comments on an earlier version of this paper. LITERATURECITED Ajilvsgi,G. 1979. Wildflowers of the Big Thicket. Texas A&M University Press, College Station, Texas. Allen, C.M. C.H. Stagg, & S.D. Parris. 1988. Analysis of the vegetation in pitcher plant bogs in two baygalls at Fort Polk in west central Louisiana. Proc. Louisiana Acad. Sci. 50:1-6. Bridges, E.L. & S.L. Orzell. 1989. Longleaf pine communities of the west gulf coastal plain. Natural Areas Journal 9:246-263. Diamond, D.D., D.H. Riskind, & S.L. Orzell. 1987. A framework for plant community classification and conservation in Texas. Texas J. Sci. 39:203-221. Fauth, J.E., J. Bernardo, M. Camara, J.W. Resetarits, J. Van Buskirk, & S.A. McCollum. 1996. Simplifying the jargon of community ecology: a conceptual approach. Amer. Naturalist 147:282-286. Folkerts,G.W. 1982. The gulf coast pitcher plant bogs. American Scientist 70:260- 267. Folkerts, G.W. 1991. A preliminary classification of pitcher plant habitats in the southeastern United States. J. Alabama Acad. Sci. 62:199-225. Harcombe, P.A., J.S. Glitzenstein, R.G. Knox, S.L. Orzell,& EL. Bridges. 1993. Vegetation of the longleaf pine region of the west gulf coastal plain. Proc. Tall Timbers Fire Ecology Conf. 18:83-104. Hermann, S.M. 1990. The ecology of pitcher plant habitats of the southeastern United States: A community profile. Unpublished report. Hermann, S.M. 1995. Status and management of Florida’s camivorous plant communities. Florida Game and Fresh Water Fish Comm. Nongame Wildl. Program Project Report. Tallahassee, Florida. Kral, R. 1955. A floristic comparison of two hillside bog localities in northeastern Texas. Field & Lab. 23:47-69. Lodwick, L.N. 1975. Net aerial primary production of three east Texas peat bogs. M.S. thesis, Baylor University, Waco, Texas. 38 PHYTOLOGIA July 1998 volume 8X 1):32-39 Louisiana Natural Heritage Program. 1988. The natural communities of Louisiana. Unpublished report, Louisiana Natural Heritage Program, Department of Wildlife and Fisheries, Baton Rouge, Louisiana. Maarel, E. van der. 1975. The Braun-Blanquet approach in perspective. Pp. 353- 359. In: R.P. McIntosh (ed.). Phytosociology. Benchmark Papers in Ecology. Dowden, Hutchinsom & Ross, Stroudsburg, Pennsylvania. MacRoberts, B.R. & M.H. MacRoberts. 1988. Floristic composition of two west Louisiana pitcher plant bogs. Phytologia 65: 184-190. MacRoberts, B.R. & M.H. MacRoberts. 1990. Vascular flora of two west Louisiana pitcher plant bogs. Phytologia 68:271-275. MacRoberts, B.R. & M.H. MacRoberts. 1991. Floristics of three bogs in western Louisiana. Phytologia. 70:135-141. MacRoberts, B.R. & M.H. MacRoberts. 1992. Floristics of four small bogs in western Louisiana with observations on species-area relationships. Phytologia 73:49-56. MacRoberts, B.R: & M.H. MacRoberts. 1993. Floristics of a bog in Vernon Parish, Louisiana, with comments on noteworthy bog plants in western Louisiana. Phytologia 75:247-258. MacRoberts, B.R. & M.H. MacRoberts. 1998a. Floristics of wetland pine savannas in the Big Thicket National Preserve, Texas. Phytologia 85(1):40-50. MacRoberts, B.R. & M.H. MacRoberts. 1998b. Floristics of muck bogs in east central Texas. Phytologia 85(1):61-72. McIntosh, R.P. 1975. Phytosociology: Benchmark Papers in Ecology, Vol. 6. Dowden, Hutchinsom & Ross, Stroudsburg, Pennsylvania. Nixon, E.S. & J.R. Ward. 1986. Floristic composition and management of east Texas pitcher plant bogs. Pp. 283-287. In: D.L. Kulhavy & R.W. Conner (eds.). Wilderness and Natural Areas in the Eastern United States: A Management Challenge. Center for Applied Studies, School of Forestry, Stephen F. Austin State University, Nacogdoches, Texas. Orzell, S.L. 1990. Texas Natural Heritage Program inventory of National Forests and National Grasslands in Texas. Unpublished report, Texas Natural Heritage Program, Texas Parks and Wildlife Department, Austin, Texas. Peet, R.K. & DJ. Allard. 1993. Longleaf pine vegetation of the southern Atlantic and eastern gulf coastal regions: a preliminary classification. Proc. Tall Timbers Fire Ecology Conf. 18:45-81. Rowell, C.M. 1949. Floral composition of a sphagnum bog in Robertson County, Texas. M.S. thesis, Texas A&M University, College Station, Texas. Sorensen, T. 1948. A method of establishing groups of equal amplitude in plant sociology based on similarity of species content. Pp. 234-249. In: R.P. McIntosh (ed.). Phytosociology. Benchmark Papers in Ecology. Dowden, Hutchinsom & Ross, Stroudsburg, Pennsylvania. Southwood, T.R.E. 1966. Ecological Methods. Methuen, London, Great Bnitain. Streng, D.R. & P.A. Harcombe. 1982. Why don’t east Texas savannas grow up into forests? Amer. Midl. Naturalist 108:278-294. Texas Natural Heritage Program (TNHP). 1993. Plant communities of Texas (Series Level). Texas Parks and Wildlife Department, Austin, Texas. Texas Organization for Endangered Species (TOES). 1992. Endangered, threatened, and watch list of natural communities of Texas. Texas Organization for Endangered Species, Austin, Texas. Turner, R.L. & J.E. Van Kley. In prep. Ecological classification system for the National Forests and surrounding areas of the West Gulf Coastal Plain. MacRoberts & MacRoberts: West Gulf Coastal Plain bog communities 39 Van Kley, J.E. In press. The vegetation of the Kisatchie sandstone hills, Louisiana. Castanea. Watson, G. 1979. Big Thicket Plant Ecology, 2nd ed. Big Thicket Museum, Saratoga, Texas. Watson, G. 1982. Plant List for the Big Thicket National Preserve. Unpublished report. Big Thicket National Preserve, Beaumont, Texas. Weakley, A.S., K.D. Patterson, S. Landall, M. Pyne, and others (compilers). 1998. International classification of ecological communities: Terrestrial vegetation of the southeastern United States. Working draft. The Nature Conservancy, Chapel Hill, North Carolina. Whittaker, R.H. 1975. Communities and Ecosystems. MacMillan, New York, New York. Wilson, J.B., I. Ullman, & P. Bannister. 1996. Do species assemblages ever recur? J. Ecology 84:471-474. Phytologia (July 1998) 85(1):40-50. FLORISTICS OF WETLAND PINE SAVANNAS IN THE BIG THICKET NATIONAL PRESERVE, SOUTHEAST TEXAS Barbara R. MacRoberts & Michael H. MacRoberts Bog Research, 740 Columbia, Shreveport, Louisiana 71104 U.S.A. & Herbarium, Museum of Life Sciences, Louisiana State University in Shreveport Shreveport, Louisiana 71115 U.S.A. ’ ABSTRACT We describe the floristics and edaphic conditions of wetland pine savannas in southeastern Texas. KEY WORDS: Wetland pine savanna, Big Thicket National Preserve, floristics, pitcher plants, Texas INTRODUCTION Southeast Texas is the western limit of once extensive wetland pine savannas (Bridges & Orzell 1989; Folkerts 1991; Harcombe et al. 1993; Marks & Harcombe 1981; Stout & Marion 1993; Streng & Harcombe 1982). In the West Gulf Coastal Plain (WGCP), wetland pine savanna is limited to coastal terraces of Allen, Calcasieu, Jefferson Davis, and Beauregard parishes in southwest Louisiana and to Jasper, Newton, Hardin, and Tyler counties in Texas (Bridges & Orzell 1989; Harcombe et al. 1993). Wetland pine savanna is open, relatively flat, and is periodically inundated. Soils are saturated during the winter and spring. There often are scattered, stunted pines; the rich herbaceous layer consists of camivorous species, grasses, sedges, and forbs. Sphagnum is abundant at some sites. Wetland pine savanna is more technically referred to as Longleaf pine-Beakrush (Pinus palustris - Rhynchospora) Series (Diamond et al. 1987), or Pinus palustris - Saturated woodland alliance (Weakley et al. 1998). MacRoberts & MacRoberts: Wetland pine savanna floristics 4) In the WGCP, wetland pine savanna is considered threatened (Texas Organization for Endangered Species 1992; Texas Natural Heritage Program 1995; Louisiana Natural Heritage Program 1993; Noss et al. 1995). There are differing opinions on the community status of wetland pine savannas in the WGCP (Bridges & Orzell 1989; Louisiana Natural Heritage Program 1988; Sheridan 1991; Texas Natural Heritage Program 1995). Do wetland pine savannas represent one or two communities? Are the northern sites, which have pitcher plants, the same or a different community from those in the south, which lack pitcher plants? Are wetland pine savannas floristically different from hillside pitcher plant bogs (Bridges & Orzell 1989; MacRoberts & MacRoberts 1993; Nixon & Ward 1986)? The purpose of this paper is twofold. First, we describe the floristics of representative wetland pine savanna in southeastern Texas. Second, using standard methods of assessing community differences, we compare two sites that have been designated as belonging to two different communities. METHODS In 1997 and 1998, we conducted a floristic inventory of two 3 ha sites on the Big Thicket National Preserve, Texas (Ajilvsgi 1979; Gunter 1993; Marks & Harcombe 1981; Watson 1979). Lance Rosier, the southernmost site, if in the Lance Rosier Unit about 8 km southwest of Kountze, Hardin County, near Little Rock Church (see Marks & Harcombe 1981 or Watson 1979 for maps of the Big Thicket National Preserve). This site is “classic” wetland pine savanna. It is about 29 km south of the natural extent of pitcher plants. The pimple mounds that occur at this site are excluded from our sample as belonging to an upland community. The second site, Turkey Creek (“Pitcher Plant Trail”), is about 32 km north of the Lance Rosier savanna in the Turkey Creek Unit about 8 km southeast of Warren, Tyler County. This site has extensive stands of pitcher plants and occurs on the southernmost edge of the extent of pitcher plants in Texas. Both sites are open. Turkey Creek has scattered shrubs and small longleaf pines. Lance Rosier has more woody vegetation, but the canopy is not more than twenty percent overall and most trees are stunted. Except for mid-winter, we conducted floristic surveys monthly and visited each site thirteen tmes between July 1997 and November 1998. All species were identified and most collected. Voucher specimens will be deposited in TEX. At each site, we established two 1? plots to measure species richness. We follow Kartesz (1994) and Jones et al. (1997) for nomenclature in most instances (but see also Nesom [1994] for Aster and Brown & Gandhi [1989] for a discussion of Hypericum; we do not distinguish between Nyssa sylvatica Marsh. and N. biflora Walt., nor between Persea borbonia (L.) Spreng. and P. palustris). The Turkey Creek site was bumed in the winter of 1997-98 and has been burned regularly for many years (Geraldine Watson, pers. comm.). The Lance Rosier site - 42 PHYTOLOGIA July 1998 volume 85 1):40-S0 was bumed in March 1983 and October 1990 (David McHugh, pers. comm.). Climatic information can be found in Marks & Harcombe (1981) and in Harcombe et al. (1993). For comparative purposes, we made brief surveys of other wetland pine savannas in both Louisiana and Texas. Soil samples were taken from the upper 15 cm and sent to A & L Laboratory, Memphis, Tennessee, for chemical analysis. RESULTS Table 1 lists the vascular plants found in the Lance Rosier and Turkey Creek study sites. Our collection number is included. Specimens without collection numbers were identified in the field. Table 1. Plants of Lance Rosier and Turkey Creek wetland pine savannas (LR = Lance Rosier, TC = Turkey Creek). SPHAGNACEAE Sphagnum sp. (LR, TC 3504] BLECHNACEAE Woodwardia areolata (L.) T. Moore [LR] W. virginica (L.) Sm. (LR, TC] LYCOPODIACEAE Lycopodiella appressa (Chapm.) R. Cranfill [LR 3734, TC 3506] L. caroliniana (L.) P. Sermolli [TC] : OSMUNDACEAE Osmunda cinnamomea L. [TC 3794] O. regalis L. [TC 3793] PINACEAE Pinus palustris P. Mill. [LR, TC] P. taeda L. [LR, TC] AMARYLLIDACEAE Hypoxis hirsuta (L.) Cov. (LR, TC 3720] BURMANNIACEAE Burmannia capitata (Walt.) Mart. [LR, TC] CYPERACEAE Carex glaucescens Ell. [LR] MacRoberts & MacRoberts: Wetland pine savanna floristics 43 Dichromena latifolia Baldw. ex Ell. [LR 3940, TC 3891] Eleocharis microcarpa Torr. (LR 3852, TC 3937] E. tuberculosa (Michx.) Roem. & Schult. [LR 3858, TC 3883] Fuirena breviseta (Coville) Coville [LR 3480] F. bushii Kral [TC 3511] Rhynchospora caduca Ell. (LR 3540] R. debilis Gale [LR 3851] R. elliotii A. Dietr. [LR 3927, TC 3948] R. filifolia A. Gray [LR 3485] . globularis (Chapm.) Small [LR 3864, TC 3934] . glomerata (L.) Vahl [LR 3646] gracilenia A. Gray [LR 3488-C, TC 3496] . inexpansa (Michx.) Vahl [LR 3541] . oligantha A. Gray [TC 3495] . plumosa Ell. [LR 3861, TC 3885] . pusilla Chapm. ex M.A. Curtis (LR 3479] . rariflora (Michx.) Ell. [LR 3867, TC 3884] Scleria georgiana Core [LR 3488, TC 3882] S. reticularis Michx. ([syn. = S. muhlenbergii Steud.] [LR 3667, TC 3499] by 30 oo WD ERIOCAULACEAE Eriocaulon decangulare L. (LR, TC] Lachnocaulon anceps (Walt.) Morong. [TC] IRIDACEAE Sisyrinchium atlanticum Bickn. [LR 3875, TC] LILIACEAE Aletris aurea Walt. [LR 3639, TC 3629] Schoenolirion croceum (Michx.) Wood [LR 3783, TC 3792] Tofieldia racemosa (Walt.) B.S.P. [TC 3513] ORCHIDACEAE Calopogon tuberosus (L.) B.S.P. [LR 3924, TC 3945] Spiranthes longilabris Lindl. [LR 4041] S. praecox (Walt.) S. Wats. [LR 3863] S. vernalis Engelm. & Gray [LR 3919] POACEAE Anthaenantia rufa (Ell.) Schultes [LR 3665, TC 3619] Aristida palustris (Chapm.) Vasey [LR 3542, TC 3618] Aristida purpurascens Poir. var. virgata (Trin.) Allred [TC 3498] Axonopus fissifolius (Raddi) Kuhlm. [LR 3653-B, TC] Coelorachis rugosa (Nutt.) Nash [LR 3637] Dichanthelium acuminatum (Swartz) Gould & Clark var. wrightianum (Scribn.) Gould & Clark [LR 3385, TC] D. consanguineum (Kunth) Gould & Clark [LR 3862] D. scabriusculum (Ell.) Gould & Clark [LR 3482, TC 3938] Eragrostis refracta (Muhl.) Scribn. [LR 3654, TC 3623] Erianthus giganteus (Walt.) Muhl. [LR] Muhlenbergia capillaris (Lam.) Trin. [LR 3656] Ad PHYTOLOGIA July 1998 Panicum brachyanthuwm Steud. (LR, TC 3634] volume 85( 1):40-S0 Panicum rigidulum Bosc. ex Nees var. pubescens (Vasey) Lelong [LR 3732, TC] Panicum tenerum Bey. ex Trin. [LR 3487, TC] Panicum verrucosum Muhl. [LR 3666, TC 3634-B] Panicum virgatum L. [LR] Paspalum floridanum Michx. [TC 3626] Paspalum laeve Michx. [LR 3663] Paspalum plicatulum Michx. [LR , TC] Paspalum praecox Walt. [LR 3731, TC 3719] Schizachyrium scoparium (Michx.) Nash [LR, TC] Tridens ambiguus (Ell.) Schultes [LR 3544, TC 3620] T. strictus (Nutt.) Nash [LR 3722] SMILACACEAE Smilax laurifolia L. [LR, TC 3632] XYRIDACEAE Xyris ambigua Bey. ex Kunth (LR, TC 3631] X. baldwiniana Schultes [TC 3879] X. caroliniana Walt. [LR, TC] X. diformis Chapm. var. diformis [LR 3657] X. louisianica Bridges & Orzell [LR 3535, TC 3501] ' X. torlaSm. [TC 3946] ACERACEAE Acer rubrum L. [LR, TC] APIACEAE Centella erecta (L.f.) Fern. [LR, TC] Eryngium integrifolium Walt. [LR, TC 3507] Oxypolis filiformis (Walt.) Britt. [LR 3478, TC] O. rigidior (L.) Raf. [TC] Ptilimnium capillaceum (Michx.) Raf. [LR, TC] AQUIFOLIACEAE Ilex coriacea (Pursh) Chapm. [LR, TC 3892] I. opaca Ait. [LR] I. vomitoria Ait. [LR, TC] ASCLEPIADACEAE Asclepias longifolia Michx. [LR 3874, TC] ASTERACEAE Arnoglossum ovatum (Walt.) H.E. Robins. [LR, TC 3508] Aster dumosus L. [LR 3648, TC 3743] Boltonia diffusa Ell. [LR 3652, TC 3509] Chaptalia tomentosa Vent. [LR 3733, TC 3624] Coreopsis linifolia Nutt. [LR 3662, TC 3627] Eupatoriwm leucolepis (DC.) Torrey & Gray [LR 3641, TC] E. rotundifolium L. (LR, TC] Helenium drummondii H. Rock [LR 3784, TC 3786] MacRoberts & MacRoberts: Wetland pine savanna floristics 45 Helianthus angustifolius L. [LR 3735] Liatris acidota Engelm. & Gray [LR 4005, TC] L. pycnostachya Michx. [LR 3648, TC] Marshallia graminifolia (Walt.) Small [LR 3378, TC 4006] Pityopsis graminifolia (Michx.) Nutt. [TC 3715] Pluchea rosea Godfrey [LR 3653, TC] CAMPANULACEAE Lobelia flaccidifolia Small [LR 3920} L. reverchonii B.L. Turner [LR, TC] CLUSIACEAE Hypericum crux-andreae (L.) Crantz [LR 3389] H. fasciculatum Lam. [LR 4001, TC 3502] H. hypericoides (L.) Crantz (LR, TC] CYRILLACEAE | Cyrilla racemiflora L. [LR, TC 3505] DROSERACEAE Drosera brevifolia Pursh [LR 3781, TC 3789] D. capillaris Poir. [LR 3860, TC 3889] FABACEAE Mimosa quadrivalvis L. var. hystricina (Small) Bameby [LR 3660] Tephrosia onobrychoides Nutt. [LR] GENTIANACEAE Bartonia verna (Michx.) Muhl. [TC 3757] Sabatia gentianoides Ell. [LR 3635, TC 4007] HALORAGIDACEAE Proserpinaca pectinata Lam. [LR 3474] HAMAMELIDACEAE Liquidambar styraciflua L. (LR, TC] LAMIACEAE Hyptis alata (Raf.) Shinners [LR 3484, TC 3625] Scutellaria integrifolia L. (LR 3876, TC 3881] LAURACEAE Persea borbonia (L.) Spreng. [LR, TC] LENTIBULARIACEAE: Pinguicula pumila Michx. [LR 3761, TC 3790] Utricularia subulata L. [LR, TC 3893] LINACEAE Linum medium (Planch.) Britt. [LR 3546, TC] 46 PHYTOLOGIA July 1998 volume 85(1):40-50 LOGANIACEAE Cynoctonum Sessilifolium (Walt.) St. Hil. [LR 3483, TC 3510] Gelsemium sempervirens (L.) St. Hil. [LR, TC] MAGNOLIACEAE Magnolia virginiana L. (LR, TC] MELASTOMATACEAE Rhexia lutea Walt. [LR 3388, TC 3944] R. mariana L. [LR 3489, TC 3942] MY RICACEAE Myrica cerifera L. (LR, TC] M. heterophylla Raf. (LR, TC] NYSSACEAE Nyssa sylvatica Marsh. [LR, TC] ONAGRACEAE Ludwigia hirtella Raf. [LR] L. linearis Walt. [LR 3538] L. pilosa Walt. (LR, TC 3500] POLY GALACEAE Polygala cruciata L. [TC 3943] P. mariana P. Mill. [LR 3726, TC] P. ramosa Ell. [LR 3877, TC] ROSACEAE Aronia arbutifolia (L.) Pers. [TC 3759] RUBIACEAE Diodia virginiana L. [LR 3475, TC] SARRACENIACEAE _ Sarraceniaalata Wood. [TC] SCROPHULARIACEAE Gratiola brevifolia Raf. [LR 3916, TC 3936] VIOLACEAE Viola lanceolata L. [LR 3762, TC 3787] V. primulifolia L. [TC 3788] There were 117 species, 76 genera, and 40 families at the Lance Rosier site. There were 106 species, 75 genera, and 41 families at the Turkey Creek site. The Sorensen Index of Similarity (Sorensen 1948) between Lance Rosier and Turkey Creek is 79, which falls into the normal range of variation for different samples of the same community type. The two | m’ plots at Turkey Creek had 20 and 24 species; the two 1 m’ plots at Lance Rosier had 19 and 21 species, indicating a rich, diverse flora. MacRoberts & MacRoberts: Wetland pine savanna floristics 47 Larry Brown (pers. comm.) informs us that Spiranthes brevilabris Lind. var. floridana (Wherry) Luer, which we did not find in out Lance Rosier study plot, occurs in the immediate area (Brown & Liggio 20418 [SBSC}). Itis rare throughout its range and is known from only a few locations in Texas (L.E. Brown pers. comm., J. Liggio pers. comm.). Table 2 gives information on soil samples from the Lance Rosier and Turkey Creek sites. Table 2. Soils of Lance Rosier and Turkey Creek study sites. eget Za Exchangeable Ions (ppm) ES Rela RN, A: MOM: ae i a ee ee ed The soils of both the Lance Rosier and Turkey Creek sites are acidic, low nutrient, and non-organic. DISCUSSION Although the main purpose of this study was to gather basic information on the floristic composition of wetland pine savannas, we were interested also in determining whether or not there was a significant difference between the floras of southern and northern sites. There has been some discussion that savannas/bogs that have pitcher plants, like Turkey Creek, are a different plant community from savannas like Lance Rosier that lack pitcher plants: the former are classified as pitcher plant bogs and the latter as wetland pine savannas (Sheridan 1991). However, the question had not been addressed quantitatively (Harcombe et al. 1993). Streng & Harcombe (1982) list the plants of a wetland pine savanna on the Hickory Creek Savanna Unit 9 km southwest of our Turkey Creek site. Even though many plants were identified only to genus, it is clear that the Hickory Creek and Turkey Creek sites are virtually identical floristically (pers. obs.). The Lance Rosier and Turkey Creek sites are not unique in the area. Near both are other wetland pine savannas with the same flora. In the vicinity of Turkey Creek are Hyatt Bog, Hickory Creek Savanna, and Kirby State Forest with its degraded wetland pine savanna. Geraldine Watson, a lifelong student of the Big Thicket, recounts many savannas that have now been destroyed. 48 PHYTOLOGIA July 1998 volume 85( 1):40-S0 The differences and similarities we have found in this study also hold for southwestern Louisiana. On the basis of incomplete surveys, the flora is essentially the same in what are called pitcher plant bogs and wetland pine savannas. In Beauregard Parish and northern Calcasieu Parish, pitcher plants occur. Farther south this species drops out of the flora, but the same basic community continues. Needless to add, for a complete understanding of the communities in the longleaf pine region of southeastern Texas and southwestern Louisiana, a much larger sample of sites from a larger part of the region would be desirable, as would additional information on hydrology, soils, geology, topographic position, importance values, cover values, and other factors that may be used to characterize sites. The fact that we found a strong similarity in flora between the Lance Rosier and Turkey Creek sites does not negate the fact that they will differ in some manner and thereby be recognizable as distinct at some level. For example, Bridges & Orzell (1989) found differences between hillside bogs and wetland pine savannas in species importance values and herbaceous species composition. We also have found such differences, not only between wetland pine savannas and hillside bogs, but among hillside bogs in different parts of the WGCP. For example, differences are apparent in Natchitoches Parish, and Vernon and Beauregard parishes. While such species as Rudbeckia scabrifolia L.E. Brown, Liatris acidota, Viola lanceolata, Dichromena latifolia, Hyptis alata, Sarracenia alata, Schoenolirion croceum, and Sabatia macrophylla Hook. are uncommon or absent in northern bogs, they are common in southern ones. These observations point to the need for thorough quantification (Harcombe et al. 1993:86). Important would be complete site specific censuses of WGCP bogs and wetland pine savannas on a north-south and east-west axis. Such concerns underline the fact that much research needs to be done on plant communities in the longleaf pine region of the WGCP and attest to the perduring problem in ecology and conservation biology of defining communities and developing community classifications (MacRoberts & MacRoberts 1998b). ACKNOWLEDGMENTS Roy Zipp, David McHugh, and Doug Hutter, Big Thicket National Preserve, and Sue Grace, National Wetlands Research Center, aided in various ways with the project. Financial support was provided, in part, by a USGS-Biological Resources Division species at risk grant to the National Wetlands Research Center. Geraldine Watson provided many hours of her time helping us see what the Big Thicket was like in years past. Joe Liggio aided with understanding soils and orchids, notably Spiranthes longilabris. Larry Brown helped with plant identifications, especially grasses, Fuirena, and brought the Spiranthes brevilabris specimen to our attention. Larry Brown, Steve Orzell, Roy Zipp, and Sue Grace reviewed an earlier version of this paper. MacRoberts & MacRoberts: Wetland pine savanna floristics 49 LITERATURE CITED Ajilvsgi,G. 1979. Wildflowers of the Big Thicket. Texas A&M University Press, College Station, Texas. Bridges, E.L. & S.L. Orzell. 1989. Longleaf pine communities of the west gulf coastal plain. Natural Areas Journal 9:246-263. Brown, L.E. & K.N. Gandhi. 1989. Notes on the flora of Texas with additions, range extensions, and one correction. Phytologia 67:394-399. Diamond, D.D., D.H. Riskind, & S.L. Orzell. 1987. A framework for plant community classification and conservation in Texas. Texas J. Sci. 39:203-221. Folkerts, G.W. 1991. A preliminary classification of pitcher plant habitats in the southeastern United States. J. Alabama Acad. Sci. 62:199-225. Gunter, P.A.Y. 1993. The Big Thicket: An Ecological Reevaluation. University of North Texas Press, Denton, Texas. Harcombe, P.A., J.S. Glitzenstein, R.G. Knox, S.L. Orzell, & E.L. Bridges. 1993. Vegetation of the longleaf pine region of the west gulf coastal plain. Proc. Tall Timbers Fire Ecology Conf. 18:83-104. Jones, S.D., J.K. Wipff, & P.M. Montgomery. 1997. Vascular Plants of Texas: A Comprehensive Checklist Including Synonymy, Bibliography, and Index. University of Texas Press, Austin, Texas. Kartesz, J.T. 1994. A Synonymized Checklist of the Vascular Flora of the United States, Canada, and Greenland. Timber Press, Portland, Oregon. Louisiana Natural Heritage Program. 1988. The natural communities of Louisiana. Unpublished report, Louisiana Natural Heritage Program, Department of Wildlife and Fisheries, Baton Rouge, Louisiana. Louisiana Natural Heritage Program. 1993. Natural plant communities in Louisiana that may be considered imperiled/cnitically imperiled. Unpublished report, Louisiana Natural Heritage Program, Department of Wildlife and Fisheries, Baton Rouge, Louisiana. MacRoberts, B.R. & M.H. MacRoberts. 1993. Floristics of a bog in Vernon Parish, Louisiana, with comments on noteworthy bog plants in western Louisiana. Phytologia 75:247-258. MacRoberts, M.H. & B.R. MacRoberts. 1998a. Wetland status of hillside bogs, wetland pine savannas, and muck bogs in the West Gulf Coastal Plain. Phytologia 85(1):22-31. MacRoberts, M.H. & B.R. MacRoberts. 1998b. Community classification of West Gulf Coastal Plain bog communities: a floristic assessment. Phytologia 85(1):32- a5: Marks, P.L. & P.A. Harcombe. 1981. Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monog. 51:287-30S. Nesom, G.L. 1994. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the New World species. Phytologia 77: 141-297. Nixon, E.S. & J.R. Ward. 1986. Floristic composition and management of east Texas pitcher plant bogs. Pp. 283-287. In: D.L. Kulhavy & R.W. Conner (eds.), Wilderness and Natural Areas in the Eastern United States: A Management Challenge. Center for Applied Studies, School of Forestry, Stephen F. Austin State University, Nacogdoches, Texas. Noss, R.F., E.T. LaRoe, & J.M. Scott. 1995. Endangered ecosystems of the United States: A preliminary assessment of loss and degradation. Biological Report 28, U.S. Departmentof the Interior, Washington, D.C. SO PHYTOLOGIA July 1998 volume 85( 1):40-50 Sheridan, P.M. 1991. What is the identity of the west gulf coast pitcher plant, Sarracenia alata Wood? Carnivorous Plant Newsletter 20: 102-110. Sorensen, T. 1948. A method of establishing groups of equal amplitude in plant sociology based on similarity of species content. Pp. 234-249. In: R.P. McIntosh (ed.). Phytosociology. Benchmark Papers in Ecology. Dowden, Hutchinsom & Ross, Stroudsburg, Pennsylvania. Stout, I.J. & W.R. Marion. 1993. Pine flatwoods and xeric pine forests of the southern (lower) coastal plain. Pp. 373-446. In: W.H. Martin, S.G. Boyce, & A.C. Echternacht (eds.). Biodiversity of the Southeastern United States. John Wiley, Inc., New York, New York. Streng, D.R. & P.A. Harcombe. 1982. Why don’t east Texas savannas grow up into forests? Amer. Midl. Naturalist 108:278-293. Texas Natural Heritage Program (TNHP). 1995. Plant communities of Texas (Series Level). Texas Parks and Wildlife Department, Austin, Texas. Texas Organization for Endangered Species (TOES). 1992. Endangered, threatened, and watch list..of natural communities of Texas. Texas Organization for Endangered Species, Austin, Texas. Watson, G. 1979. Big Thicket Plant Ecology, 2nd ed. Big Thicket Museum, Saratoga, Texas. Weakley, A.S., K.D. Patterson, S. Landall, M. Pyne, and others (compilers). 1998. International classification of ecological communities: Terrestrial vegetation of the southeastern United States. Working draft. The Nature Conservancy, Chapel Hill, North Carolina. Phytologia (July 1998) 85(1):$1-60. WOODY VEGETATION DIVERSITY OF LONGLEAF PINE COMMUNITIES IN CALCASIEU PARISH, LOUISIANA Ray Neyland, Harry A. Meyer, & Heather Harrington Department of Biological and Environmental Sciences, McNeese State University, Lake Charles, Louisiana 70609-2000 U.S.A. ABSTRACT Located in southwestern Louisiana, Calcasieu Parish once supported large areas of undisturbed longleaf pinelands. These pinelands were clear cut around 1900 and have been converted largely to forest, range, and cropland. Although virgin longleaf pinelands no longer exist in the Parish, a few remnant sites retain many of their presettlement plant species and characteristics. Three distinctly different longleaf pine communities were identified and sampled to determine woody species diversity and importance. Diversity values of the three communities differed significantly from each other. The acid longleaf pine flatwoods community was less diverse than either the hillside seepage bog community or the sodic longleaf pine flatwoods community. The low diversity value for the acid longleaf pine flatwoods community was due primarily to low species richness when compared to the other two communities. Although the hillside seepage bog and the sodic longleaf pine flatwoods each had identical species richness values, the disparity in their diversity values is explained primarily by the greater evenness of species abundance at the sodic longleaf pine flatwoods community. The various mixtures of woody species sampled suggest a high degree of heterogeneity among the sites chosen for study. For example, of the eleven different species found among the three communities, only Pinus palustris was present at all three. Pinus palustris was the most important in all but the sodic longleaf pine flatwoods community where it was second to P. elliotii. The importance value for all combined communities was clearly dominated by P. palustris at 140.69. Pinus elliottii had the second highest importance value for all combined communities at 62.83 with Myrica cerifera third at 31.58. Relative coverage for all communities combined was also dominated by P. palustris at 55.82% with P. elliottii and M. cerifera at 31.73% and 6.56% respectively. KEY WORDS: Calcasieu Parish, Louisiana, longleaf pine communities, woodland remnants a1 ‘32 PHYTOLOGIA July 1998 volume 85 1):51-60 The longleaf pineland region of the West Gulf Coastal Plain (Bridges & Orzell 1989) extends across northern Calcasieu Parish (Figure 1). This region onginally extended from southeastern Texas to north-central Louisiana (Wahlenberg 1946; Little 1971). These pinelands were clear cut in the early 1900's (Roy & Midkiff 1988; Smith 1991) and now exist primarily as forest, range and cropland (Roy & Midkiff 1988). Remnant longleaf pine communities of the West Gulf Coastal Plain identified by Orzell (1987) and Bridges (1988) are currently threatened by grazing, silvaculture, and wildlife management (Bridges & Orzell 1989). The once virgin longleaf pinelands as described by Schwarz (1907) and Wahlenberg (1946) are now rare in Louisiana (Smith 1991). However, a few comparatively undisturbed sites within Calcasieu Parish appear to retain many of their presettlement plant species and remain recognizable longleaf pine communities (sensu Bndges & Orzell 1989; Smith 1991, 1996). These rare sites are small and may be considered remnants of a once dominant ecosystem. The purpose of this study is to characterize the adult woody vegetation of three remnant longleaf pine communities in Calcasieu Parish in terms of density, frequency coverage, and importance. These communities are referred to as hillside seepage bogs, sodic longleaf pine flatwoods (sodic flatwoods), and acid longleaf pine flatwoods (acidic flatwoods). This terminology is sensu Smith (1996). STUDY AREA Located in southwestern Louisiana, Calcasieu Parish is adjacent to Beauregard Parish to the north, Jefferson Davis Parish to the east, Cameron Parish to the south, and the State of Texas to the west (Figure 1). Approximately 46% of the Parish's land is devoted to either agriculture or rangeland; approximately 23% is woodland; 11% is marsh; 4% swamp; and the remaining area is urban (Roy & Midkiff 1988). Land use is devoted primarily to timber, cattle, rice, and soybeans. Petroleum-related industries are present in the Parish and are concentrated around the area of Westlake. Calcasieu Parish is noteworthy for the abrupt transition from the longleaf pinelands in the north to the prairie and coastal marshes in the south (Figure 1). The first study site is a hillside seepage bog situated about 8 km northeast of DeQuincy; (93° 23’ 83” x 30° 28’ 92”); range and township (RIOW x T7S Sect. 1) (Figure 1). This is the only known representative of this community type in the Parish. The site extends approximately 600 m on a north-south-axis by 36 m on an east-west axis and is bounded to the west by a semi-evergreen broadleaf acid seep forest (sensu Bridges & Orzell 1989), and to the east by a stand of Pinus elliotii Engelm. This bog appears to be the headwaters of a small unnamed stream. The soil type is “Glenmora silt loam” (Roy & Midkiff 1988) and is characterized by an extremely acidic silt loam surface layer and a strongly acid silty clay loam subsoil. Distinctive herbaceous species include Rhynchospora oligantha A. Gray, Sarracenia alata (Wood) Wood, and Utricularia cornuta Michx. Charred bases of P. palustris Mill. indicate the site had bummed within the past several years. Neyland etal.: Woody vegetation in Calcasieu Parish 53 Coastal Marsh Bottomland Forests Coastal Prairie Swampland Longleaf Pine Figure 1. Area of longleaf pineland in relation to other regions within Calcasieu Parish, Louisiana. Inset shows the location of the Parish within the State. Study site #1 = hillside seepage bog; #2 = sodic flatwoods; #3 = acidic flatwoods. 54 PHYTOLOGIA July 1998 volume 85(1):51-60 The second study site is a sodic flatwoods that is situated near a few other similar sites in the western part of Calcasieu's longleaf pineland region (Figure 1). These flatwoods are typically saturated during the winter and spring but may become very dry during summer droughts. The specific site studied is situated about 22 km southwest of DeQuincy; (93° 34’ 15” x 30° 18’ 74”); range and township (R11W x T8S Sect. 35) (Figure 1). Itis bounded to the south by the Houston River and to the west by Creek Road. The community is approximately 2.5 km long on its east-west axis and 2.0 km long on its north-south axis. Distinctive herbaceous species include Spartina spartinae (Trin.) Hitch., Chaetopappa asteroides (Nutt.) DC., Evolvulus sericeus Sw., and Liatris punctata Hook. The understory woody vegetation is. stunted. The soil type is “Brimstone” and has a very strongly acidic silt loam layer, a subsurface that is medium acid silt loam, and a subsoil of alkaline silty clay that is high in sodium (Roy & Midkiff 1988). The third study site is an acidic flatwoods and is situated in the central part of the longleaf pineland region, about 10 km southeast of DeQuincy; (93° 20’ 85” x 30° 22’ 63”); range and township (ROW x T7S Sect. 11) (Figure 1). This acidic flatwoods extends about 1.2:km on its north-south axis and about 0.4 km on its east-west axis and is bounded on the east by Holbrook Park Rd. and to the west by cleared land. The soil type is “Caddo-Messer silt loam” and has a very strongly acidic surface and subsurface layer (Roy & Midkiff 1988). The subsoil is strongly acid silty clay loam or silt loam. The terrain is level to gently rolling with small elevated areas termed “pimple mounds” by Holland ef al. (1952). Under natural conditions and frequent fires, these flatwoods support a sparse canopy of longleaf pine with few other tree species (Bridges & Orzell 1989). The planting of pine plantations and fire suppression have nearly eliminated these communities in the Parish. Distinctive herbaceous species in this community include Stylisma aquatica (Walter) Raf. and Platanthera nivea (Nutt.) Luer. To our knowledge, this is the only acidic flatwoods community in Calcasieu Parish that supports a rich herbaceous layer. MATERIALS AND METHODS The three representative community types chosen for this study share the following characteristics: 1) longleaf pine is a major component; 2) community maintenance requires frequent fires; 3) herbaceous ground cover is species rich. Each site was selected to represent one of the three longleaf pine communities in Calcasieu Parish that shares these criteria. Although these sites are not pristine, we consider them recognizable longleaf pine communities (sensu Bridges & Orzell 1989; Smith 1991, 1996). Each sampled area measured 100 m along its long axis and 36 m on its short axis. Data were collected on tree and shrub species (vines were excluded) with a diameter at breast height (DBH); (1.3 m) of 1 cmor greater. Randomly selected ten by ten meter quadrats (plots) were used for sampling within the defined study sites. Plot size was based on that recommended for forest trees by Cox (1996) and Brower et al. (1998). Neyland etal.: Woody vegetation in Calcasieu Parish 55 The number of plots sampled per site was determined by constructing a species area curve to ensure we encountered the bulk of species present (Cox 1996). Specifically, a plot of cumulative number of species encountered versus cumulative number of plots sampled indicated fifteen plots was adequate to sample nearly all of the species present. Sampling for hillside seepage bog, sodic flatwoods and acid flatwoods sites were performed on April 8, May 3, and May 20, 1998 respectively. Nomenclature follows Kartesz (1994). Frequency, density, coverage (basal area) and importance values were calculated to determine the dominant woody species (Cox 1996). Basal area was estimated from DBH values. The importance value is equal to the sum of relative frequency, relative density, and relative coverage. These values were calculated for each of the three sites individually and for all sites combined. Species diversity was calculated using Simpson's Diversity Index (Cox 1996) and was calculated from density data. Because we consider the few small “pimple mounds” to be “intrusions” into the acidic flatwoods site, woody species were excluded from sampling. However, a complete inventory of the woody species inhabiting these mounds was made and is included in the following section. RESULTS Species area curves (Cox 1996) suggest that fifteen samples were sufficient to encounter the bulk of species present at each site. That is, the final plot number for each site was positioned well into the asymptotic portion of the curve. Woody vegetation diversity values among the communities differed significantly (p < 0.05) (Table 1). The low diversity value for the acidic flatwoods site was due primarily to low species richness as compared to the other two communities. Although the hillside seepage bog and the sodic flatwoods site each had identical species richness values (Table 1), the disparity in diversity values is explained primarily by greater evenness of species abundance at the sodic flatwoods. Density, frequency, coverage, and importance values for all sites are summarized in Table 2. The various mixtures of woody species sampled suggest a high degree of heterogeneity among the three sites. Of the eleven different species discovered among the sites, only Pinus palustris was present in all three. Myrica cerifera L. was present in all three communities but no individuals met our size criterion for sampling in the acidic flatwoods due to a recent burn. Additionally, seven species were present in only one of the three sites: P. elliottii, Ilex vomitoria Aiton, Quercus marilandica Muench, Viburnum dentatum L., I. coriacea (Pursh) Chapm., Liquidambar styraciflua L., and Sapium sebiferum (L.) Roxb. (Table 2). Heterogeneity among the communities is also demonstrated by the importance values of P. palustris. That is, although P. palustris was the most important species for all communities combined (Table 3), it was only the second most important species in the sodic flatwoods site (Table 2). 56 PHYTOLOGIA July 1998 volume 85(1):51-60 Table 1. Simpson's Diversity Index, variance, and species richness of woody vegetation by study site. pels scarey take doinat ts Index (Ds ) Variance (S2) Species Richness oS a ORE Hillside Bog 0.495 EE Cell ho 0008 0.165 Table 2. Density, frequency, coverage and importance values by species by individual site. Species are arranged in order of importance. Density | Relative | Frequency | Relative | Coverage | Relative} Importan pecies (#/plot) | density Frequenc wae coverage} Value ea toe tee ogc noth aie das and oa < Vey on iad Je bounded on 'e Git br Tes bead oe 7 op ho a PieMilisdeBog escent a perpen | os oon a Sic a Ma a Da Nyse biflora 0781138 [oad fr. 33.59 5.85 Sabian ebifenan> [007 +108 [007 1-300} 021- L004 ed Aol ee ee loves fall fms ie pooore|\ wee [ait One fm aero Se eich ef peeps ey ee ee | ae a a a 38.91 29. Ss = $s 48.12 | 116.68 32.80 38.75 | _ 86.15 11.55 [0.60 [26.85 [ 44.22 [3.15 41.25 12.35 14.60 36.08 0.46 | 7.94 ee ee ee at ee ree cee ADSI Cee SNE MN eT Rs a SE ES RN OS 90.07 85.11 99.17 | 274.35 Re et a 8 T O78 T1320 | I EE RB Neyland et al.: Woody vegetation in Calcasieu Parish 7 Table3. Density, frequency, coverage, and importance values by species summarized for all sites. Species are arranged in order of importance. pecies Density | Relative | Frequency | Relative | Coverage | Relative | Importance (#/plot) | Density Frequency | (¢m2/piot)| Coverage | Value © i i Pa GE DS Se ee Sea 51.17] 0.62] 33.70] 398.49| 55.82] 140.60 (Pinus elliot | 0.98] 19.14] _0.22| _11.96| 226.55] 31.73] 62.83 [Myrica ceriera______| 0.53] 10.35| 0.27] 14.67] 46.82] 6.56] 31.58 Mex vomitoria | 0.29] _ 5.66| 0.20] 10.87] 14.81] 2.07| 18.05] Mex coriacea | 0.24, 4.69| 0.18] 9.78] 10.91] 1.53] 16.00] [Nyssa biflora | 0.27| 5.27| 0.16] 870] 11.19] 1.57| 15.54) [Quercus marilandica __| 0.07| 1.37] 0.07] 3.80] 1.86] 0.26| 5.43] Viburnum deniatum | 0.04[ 0.78] 0.04] 2.17] 2.18] 0.31] 3.26) quidambar styracifiua | 0.04] 0.78] 0.04] 2.17] 0.59] 0.08] 3.03, (Sapium sebijerum___| 0.02| 039] 0.02] 1.09] 0.07 Importance for all combined communities was dominated by Pinus palustris at 140.69 (Table 3). Pinus palustris was the most important in all but the sodic flatwoods community where it was second to P. elliottii (Table 3). Pinus ellionii had the second highest importance value for all combined communities at 62.83 with Myrica cerifera third at 31.58 (Table 3). Relative coverage for all communities combined was dominated by P. palustris at 55.82% and P. elliottii at 31.73% (Table > Woody species that occurred in quadrats of the hillside seepage bog but did not meet the minimum size requirements for sampling included Acer rubrum L., Aronia arbutifolia (L.) Elliott, Hypericum brachyphyllum (Spach) Steud., and Magnolia virginiana L. Because neither Acer rubrum nor M. virginiana had sufficient time for regrowth since a recent burn, no individuals met our criterion for sampling. Additionally, these two species are rare in this particular community. Although H. brachyphyllum and Aronia arbutifolia are common at this site, both are small shrubs and no individuals met our sampling criterion. The woody vine, Smilax laurifolia L.., was also present. Rare individuals of Magnolia grandiflom L. were observed in the sodic flatwoods site but were not included in any sampled quadrats. Although the woody shrub Ilex opaca Aiton was common in the site, no individuals met our size requirements for sampling. Woody vines present included Smilax smallii Morong and Berchemia scandens (Hill) K. Koch. . Woody species that occurred in the acidic flatwoods quadrats but did not meet the minimum size requirements for sampling included Baccharis halimifolia L., Sapium sebiferum, and Myrica cerifera. Because of a controlled burn in the fall of 1996, no individuals of these three species were sufficiently large enough to meet the sampling 58 PHY TOLOGIA July 1998 volume 85 1):51-60 criterion. Of these three species encountered, only M. cerifera appeared to be common on the site. The woody vine, Smilax laurifolia, was also present. A complete inventory of the woody species inhabiting the pimple mounds within the acidic flatwoods community includes Sapium sebiferum, Acer rubrum, Myrica cerifera, Sassafras albidum (Nutt.) Nees, Quercus falcata Michx., Q. nigra L., Ilex opaca, and Liquidambar styraciflua. DISCUSSION Although not pristine, the three remnant sites analyzed in this study resemble longleaf pine communities before settlement and clear cutting. However, the current state of these pinelands is undoubtedly different from the time prior to settlement. For example, the most important species found in the sodic flatwoods site, Pinus elliottii, is native only in Louisiana parishes east of the Mississippi River (Flora of North America Committee 1993) and, therefore, would have been absent in presettlement times. Although this commercially introduced species historically has been used for its naval stores, P. elliottii has become an increasingly important plantation pine for lumber and plywood (Flora of North America Committee 1993). The establishment of P. elliottii has almost certainly displaced other native woody species. Steps have recently been taken to preserve at least a part of the sodic flatwoods in Calcasieu Parish. The Nature Conservancy of Louisiana has recently purchased approximately 240 contiguous acres of sodic flatwoods along Persimmon Gully that are adjacent to study site number two (Richard Martin, pers. comm., 1998). The acid flatwoods and hillside seepage bog sites support a few individuals of the introduced Sapium sebiferum. This native of subtropical China, commonly called Chinese tallow, has been planted in several areas of the world as an ornamental and for the production of vegetable tallow and stillingia oil from its fruits (Kahn et al. 1973). This species was introduced to the Southeast by the U.S.D.A. Bureau of Plant Industry in the early 1900's to establish local soap industries (Jamieson & McKinney 1938). The ability of Sapium to aggressively invade and quickly alter habitat is well documented (Cameron & Spencer 1989; Bruce et al. 1995; Neyland & Meyer 1997). At present, the incidence of Sapium at these two sites is low. However, it remains an Open question to what extent this species will alter the ecology of these communities. Additionally, species composition in the hillside seepage bog has been altered since settlement times by the introduced Pinus elliottii. Specifically, all but the lowermost 36 m of the hillside seepage bog now supports a dense stand of P. elliottii. The future of these remnant longleaf pine communities in Calcasieu Parish is uncertain. Most remain threatened by urbanization, wildlife management practices, and silvaculture. Without active protection, these rare communities may be destined for extinction in Calcasieu Parish. Neyland etal.: Woody vegetation in Calcasieu Parish 59 ACKNOWLEDGMENTS We thank Latimore Smith from the National Heritage Program of the Louisiana Department of Wildlife & Fisheries for reviewing previous drafts of this paper and for providing us the location of the hillside seepage bog study site. We also thank Lowell E. Urbatsch (Louisiana State University) and Mark Paulissen (McNeese State University) for reviewing this paper prior to submission. LITERATURECITED Bridges, E.L. 1988. A preliminary survey for potential natural areas in the pine flatwoods region of southwestern Louisiana. Unpublished report for the Louisiana Natural Heritage Program, Baton Rouge, Louisiana. Bridges, E.L. & S.L. Orzell. 1989. Longleaf pine communities of the West Gulf Coastal Plain. Natural Areas Journal 9:246-263. Brower, J.E. , J.H. Zahr, & C.N. von Ende. 1998. Field and Laboratory Methods for General Ecology, Fourth Edition. Wm. C. Brown, Dubuque, Iowa. Bruce, K.A., G.N. Cameron, & P.A. Harcombe. 1995. Initiation of a new woodland type on the Texas Coastal Prairie by the Chinese tallow tree (Sapium sebiferum (L.) Roxb.). Bull. Torrey Bot. Club 122:215-225. Cameron, G.N. & R. Spencer. 1989. Rapid leaf decay and nutrient release in a Chinese tallow forest. Oecologia 80:222-228. Cox, G.W. 1996. Laboratory Manual of General Ecology, Sixth Edition. William C. Brown, San Diego State University. Flora of North AmericaCommittee. 1993. Floraof North America North of Mexico, Vol. 2. Pteridophytes and Gymnosperms. Oxford University Press, New York, New York. Holland, W.C., W. Hough, & G.E. Murry. 1952. Geology of Beauregard and Allen Parishes. Louisiana Geological Survey, Baton Rouge, Louisiana, Geological Bulletin 27. Jamieson, G.S. & R.S. McKinney. 1938. Stillingia Oil. Oil and Soap. 15:295-296. Kahn, F.W., K. Kahn, & M.N. Malik. 1973. Vegetable tallow and stillingia oil from the fruits of Sapium sebiferum, Roxb. Pakistan Journal of Forestry 23:257-266. Kartesz, J.T. 1994. A Synonymized Checklist of the Vascular Flora of the United States, Canada, and Greenland. 2nd. ed. Timber Press, Portland, Oregon. Little, E.L. 1971. Atlas of United States Trees: Volume 1- conifers and important hardwoods. USDA Forest Service Miscellaneous Publication 1146, Washington, By. Neyland, R. & H.A. Meyer. 1997. Species diversity of Louisiana chenier woody vegetation remnants. J. Torrey Bot. Soc. 124:254-261. Orzell, S.L. 1987. Interim status report on natural heritage program inventory of national forest lands. Report submitted to U.S. Forest Service, Lufkin, Texas. Roy, A.J. & C.T. Midkiff. 1988. Soil survey of Calcasieu Parish, Louisiana. USDA, Soil Conservation Service. Schwarz, G.F. 1907. The Longleaf Pine in Virgin Forest. John Wiley & Sons, New York, New York. 60 PHYTOLOGIA July 1998 volume 85( 1):51-60 Smith, L.M. 1991. Louisiana longleaf, an endangered legacy. Louisiana Conservationist 43:24-27. Smith, L.M. 1996. The rare and sensitive natural wetland plant communities of interior Louisiana. Louisiana Department of Wildlife and Fisheries, Natural Heritage Program, Baton Rouge, Louisiana. Wahlenberg, W.G. 1946. Longleaf pine: its use, ecology, regeneration, protection, growth, and management. Charles Lathrop Pack Forestry Foundation, Washington, D.C. Phytologia (July 1998) 85(1):61-73. FLORISTICS OF MUCK BOGS IN EAST CENTRAL TEXAS Barbara R. MacRoberts & Michael H. MacRoberts Bog Research, 740 Columbia, Shreveport, Louisiana 71104 U.S.A. & Herbarium, Museum of Life Sciences, Louisiana State University in Shreveport, Shreveport, Louisiana 71115 U.S.A. ABSTRACT We describe the floristics and edaphic conditions of muck bogs in the post oak savanna region of east central Texas. These bogs are floristically different from east Texas hillside seepage bogs. KEY WORDS: bog, Gus Engeling Wildlife Management Area, Sarracenia, post oak savanna, pitcher plant, floristics, Texas INTRODUCTION Recent studies of pitcher plant habitat (bogs) in the West Gulf Coastal Plain (WGCP) have been confined to the piney woods region of east Texas and western Louisiana (Nixon & Ward 1986; Allen et al. 1988; MacRoberts & MacRoberts 1988, 1990, 1991, 1992, 1993; Bridges & Orzell 1989a; Harcombe et al. 1993). Studies of muck bogs in the post oak savanna region of east central Texas (Rowell 1949a, 1949b; Kral 1955; Lodwick 1975; Starbuck 1984; Bridges & Orzell 1989b) show a type of bog that differs in many respects from more eastern bogs. Among other things, these bogs quake and have a very high organic content and biomass. Analyses of peat deposits at some of these sites (Potzger & Tharp 1943, 1947, 1954; Graham & Heimsch 1960; Larsen et al. 1972; Bryant 1977) indicate that bogs with high organic content have been present in east central Texas for thousands of years. Kral (1955), Rowell (1949a, 1949b), and Lodwick (1975) provide a general description of muck bogs. The bog edge, often up to a meter higher than the bog center, is sandy and has low organic content. During short droughts, the edge may temporarily become dry. The bog flattens toward its center. The soil is completely saturated throughout the year and consists of a thick organic slurry that is kept wet by seepage from surrounding xeric sandhills of higher elevation. The bogs are firm enough to walk on and consist of masses of intertwined roots and rhizomes. The 61 62 PHYTOLOGIA July 1998 volume 8S 1):61-73 mass of vegetation grows mainly on root hummocks, although some species are floating or anchored in the slurry amongst the hummocks. Sphagnum is present and buildup of organic matter is high. Peat depths of over five meters have been measured. In places, the surface quakes, and it is possible to shove a pole two to three meters down with little resistance. Muck bogs are open and generally treeless, and appear to develop on blocked meander streams. Shrub thickets may develop when fire is excluded. Many muck bogs have pitcher plants and a rich herbaceous layer with an array of carnivorous species, grasses, sedges, and xyrids (Folkerts 1991; Kral 1955; Lodwick 1975; Rowell 1949a, 1949b; Starbuck 1984; Bridges & Orzell 1989b). In the center of the bog, there often is a stream. The purpose of this paper is to describe muck bogs at Gus Engeling Wildlife Management Area (GEWMA), Anderson County, Texas. It was of interest to us to extend our research on bog communities of the West Gulf Coastal Plain to these post oak savanna bogs and associated marshes. They have been little studied, and most of what has been recorded about them is unpublished (Lodwick 1975; Rowell 1949a; Starbuck 1984). STUDY SITES/METHODS In 1997 and 1998, we conducted a systematic study of the flonistics of two muck bogs on the 4436 ha GEWMA, Anderson County, in the post oak savanna region of east central Texas (Telfair 1988; Lodwick 1975; Johnson 1986). The area is gently rolling to hilly and is drained by Catfish Creek, a tributary of the Trinity River. Most of the streams are spring fed and flow year round. The upland soils are rapidly permeable sands (Telfair et al. n.d.; Hauke & Rose n.d.). Detailed floristics were done on two bogs: Chester’s Bog is on Gibson Branch at the northern end of GEWMA and Andrew’s Bog is east of DD Spring, also at the northern end of GEWMA. The two bogs are about 1 km apart. Chester’s Bog is down stream from an artificial lake, which has undoubtedly affected its history by altering its hydrology. Andrew’s Bog is not affected by any artificial impoundment or any man-altered drainage. Surveys were conducted on a monthly basis, except for the midwinter months. From July 1997 to July 1998, each bog was surveyed ten times. The northern edge of Andrew’s Bog was surveyed at two sites totaling about3 ha. An area of about 2 ha of Chester’s Bog was surveyed. We follow Kartesz (1994) for nomenclature in most cases, but see Jones et al. (1997) and Nesom (1994). Soil samples were analyzed by A & L Laboratories, Memphis, Tennessee. Because the organic matter was so high, we ran a “manure” analysis on one sample to compare results (see A & L Analytical Services List for procedures). MacRoberts & MacRoberts: Floristics of muck bogs 63 The early fire history of these bogs is unknown. Chester’s Bog was bumed in early 1990; the precise month is not known. It was badly overgrown with shrubs and was hand cleared in the summer of 1995. Andrew’s Bog was bumed sometime in 1980 and in February 1985, 1994, 1996, and 1998. For east Texas, precipitation is generally uniformly distributed throughout the year, averaging about 100 cm. Summers are long and hot; temperatures rise to 35° C, which, combined with short droughts, translates into dry conditions with streams sometimes drying up: 1998 was a drought year. Winters are mild with few days of freezing weather (typically about 268 frost-free days). Mean annual temperature is about 20° C, with an average January temperature of 7° C and average July temperature of 27° C. RESULTS Table 1 lists the vascular plants found in Chester’s and Andrew’s bogs. (A = present in Andrew’s Bog, C = present in Chester’s Bog: number in brackets is our collection number. No number indicates no collection. Specimens will be deposited at VDB-BRIT and TEX.) Table 1. Plants of Chester’s (C) and Andrew’s (A) bogs. Numbers refer to MacRoberts & MacRoberts collection numbers. SPHAGNACEAE Sphagnum [A, C] BLECHNACEAE Woodwardia areolata (L.) T. Moore [A, C 358]] W. virginica (L.) Sm. [A] LYCOPODIACEAE Lycopodiella appressa (Chapm.) Cranfill [A 3979, C 3694] OSMUNDACEAE Osmunda cinnamomea L. [A, C] O. regalis L. [A, C] SELAGINELLACEAE. Selaginella apoda (L.) Fern. [C 3775] AMARYLLIDACEAE Hypoxis hirsuta (L.) Cov. [A 3819, C] 64 PHYTOLOGIA July 1998 volume 85(1):61-73 ARACEAE Peltandra virginica (L.) Schott [A 4035] BURMANNIACEAE Burmannia capitata (Walt.) Mart. [A, C 3563] CYPERACEAE Carex atlantica Bailey [A 3814, C 3841] Carex glaucescens Ell. [C] Carex lurida Wahlenb. [A 3815, C 3842] Carex stricta Lam. [A 3813, C 3770] Cladium mariscoides [A 3990] Cladium mariscus (L.) Pohl subsp. jamaicense (Crantz) Kukenth. [C 3993] Cyperus odoratus L. [C 3690] Cyperus haspan L. [C 3436] Eleocharis equisetoides (Ell.) Torr. [A 3952, C 3961] E. olivacea Torr. [C 3708] E. tortilis (Link) Schultes [A 3802, C 3846] Fuirena squarrosa Michx. [A 3602, C 3434] Rhynchospora cephalantha A. Gray [C 3561] R. chalarocephala Fern. & Gale [A 3992, C 3999-A] R. corniculata (Lam.) A. Gray [A 3981, C] R. globularis (Chapm.) Small [A 3834, C] R. glomerata (L.) Vahl [(C 3411] R. gracilenta A. Gray [A 3595, C] R. rariflora (Michx.) Ell. [A 3832, C] Scirpus cyperinus (L.) Kunth [A, C 4026] Scleria reticularis Michx. [A 3599, C 3558] ERIOCAULACEAE Eriocaulon decangulare L. [A 3826, C 3410] E. kornickianum van Heurck & Muell.-Arg. [A 3950] IRIDACEAE Iris virginica L. [A 3821, C 3849] Sisyrinchium langloisii E. Greene [A 3805] JUNCACEAE Juncus coriaceus Mack. [C 3584] J. diffusissimus Buckl. [A, C 3972] J. effusus L. [A 3810, C 3838] J. marginatus Rostk. [A 3833, C 3996] J. scirpoides Lam. [A] J. trigonocarpus Steud. [A 3603, C 3568] LILIACEAE Zigadenus densus (Desr.) Fern. [A 3820] MAYACACEAE Mayaca fluviatilis Aubl. [A 3673, C 3579] MacRoberts & MacRoberts: Floristics of muck bogs F 65 ORCHIDACEAE Calopogon tuberosus (L.) B.S.P. [A, C 3974] Platanthera ciliaris (L.) Lindl. [C 3444] Pogonia ophioglossoides (L.) Ker.-Gawl. [A 3822, C 3837] Spiranthes cernua (L.) L.C. Rich. [A 3682, C 3684] POACEAE Agrostis hyemalis (Walt.) B.S.P. [A 3801] Agrostis scabra Willd. [A 3741, C 3697] Andropogon glomeratus (Walt.) B.S.P. [A 3600, C 3555] Chasmanthium laxum (L.) Yates [C 3437] Cinna arundinacea L. [C 3583-B] Coelorachis rugosa (Nutt.) Nash [A 368/] Dichanthelium dichotomum (L.) Gould var. dichotomum [A, C 3839] D. scabriusculum (Ell.) Gould & Clark [A 3816, C 3711} D. scoparium (Lam.) Gould [A 3975, C] Erianthus giganteus (Walt.) Muhl. [A 3596, C 3550] Leersia hexandra Sw. [C 3696] L. oryzoides (L.) Sw. [C 3693] Panicum rigidulum Bosc. ex Nees [A 3601, C 3553] Panicum verrucosum Muhl. [A, C 3673] Panicum virgatum L. [A, C 3593] Paspalum plicatulum Michx. [C 3582] Paspalum praecox Walt. [A 3678] Sacciolepis striata (L.) Nash [A] SMILACACEAE Smilax glauca Wall. [A, C] SPARGANIACEAE Sparganium americanum Nutt. (C 3995] XYRIDACEAE Xyris ambigua Bey. ex Kunth [A, C 3401]: X. baldwiniana Schultes [A 3951, C 3962] X. diformis Chapm. var. diformis [A, C 3407] X. jupicai L.C. Rich. [C 3409] X. toriaSm. [A 3985, C 3403] ACERACEAE Acer rubrum L. [A, C 4028] APIACEAE Centella erecta (L.f.) Fern. [A, C] Eryngium integrifolium Walt. [A, C 3422] Hydrocotyle umbellata L. [A, C] Oxypolis rigidior (L.) Raf. [C 3571] Ptilimnium capillaceum (Michx.) Raf. [A, C 3968] -P. costatum (Ell.) Raf. [A, C 3412] 66 PHYTOLOGIA July 1998 volume 85( 1):61-73 ASCLEPIADACEAE Asclepias rubra L. [A, C 3417] ASTERACEAE Aster lateriflorus (L.) Britt. [A 3680, C] A. puniceus L. var. scabricaulis (Shinners) A.G. Jones [A 3672, C 3710] A. umbellatus P. Mill. (C 3713] Boltonia diffusa Ell. [LR 3604, C] Eupatorium fistulosum Barratt [C] E. perfoliatum L. [A, C] E. rotundifolium L. [A, C 3415] Helianthus angustifolius L. [A, C 3548] Liatris pycnostachya Michx. [C] Mikania scandens (L.) Willd. [C 3585] Pluchea rosea Godfrey [A, C] Solidago rugosa P. Mill. [A 3598, C 3569] Vernonia missurica Raf. [A, C 3414] BETULACEAE Alnus serrulata (Ait.) Willd. [A, C 4030] CAMPANULACEAE Lobelia puberula Michx. [A 3597, C 3572] L. reverchonii B.L. Turner [A 3683, C 3704]] CAPRIFOLIACEAE Viburnum nudum L. [A, C 3419] CLUSIACEAE Hypericum crux-andreae (L.) Crantz [A 4036, C] H. mutilum L. [A, C 3441] Triadenum virginicum (L.) Raf. [A, C 3554] DROSERACEAE Drosera brevifolia Pursh [A 3829, C] D. capillaris Poir. [A, C 3576] ERICACEAE Rhododendron oblongifolium (Small) Millais [C 3973] FABACEAE Apios americana Medic. [A] GENTIANACEAE Bartonia paniculata (Michx.) Muhl. [A 3679, C 3688] HY DROPHYLLACEAE Hydrolea ovata Choisy [A 4037] MacRoberts & MacRoberts: Flonstics of muck bogs LAMIACEAE Lycopus rubellus Moench [A, C 3586] Scutellaria integrifolia L. [A 3807, C 3848] LENTIBULARIACEAE Utricularia cornuta Michx. [A 3808, C] U. gibba L. [A, C 3580] U. juncea Vahl. [A 3988, C 3994] U. subulata L. [A 3828, C 3847] LINACEAE Linum striatum Walt. [C 3424] MALVACEAE Hibiscus moscheutos L. subsp. lasiocarpus (Cav.) O.J. Blanchard [A 3984] MELASTOMATACEAE Rhexia mariana L. [A, C 3578] R. virginica L. [A, C 3565] MYRICACEAE Myrica cerifera L. [A, C 3421] NYSSACEAE Nyssa sylvatica Marsh. [A, C] ONAGRACEAE Ludwigia alternifolia L. [A 3980, C 3566] L. decurrens Walt. [C 3687] L. glandulosa Walt. [A, C] POLY GALACEAE Polygala cruciata L. [A, C 3420] P. sanguinea L. [A 3954, C] RUBIACEAE Cephalanthus occidentalis L. [C] SALICACEAE Salix nigra Marsh. (C] SARRACENIACEAE Sarracenia alata Wood. [A 3674, C] SAURURACEAE . Saururus cernuus L. [A, C 3425] SCROPHULARIACEAE Agalinis fasciculata (Ell.) Raf. [A, C 369/] Gratiola brevifolia Raf. [A 3976, C] 67 68 PHY TOLOGIA July 1998 volume 85( 1):61-73 URTICACEAE Boehmeria cylindrica (L.) Sw. [A, C 3559] VALERIANACEAE Valerianella woodsiana (Torr. & Gray) Walp. [A 4034] VIOLACEAE Viola primulifolia L. [A 3812, C 3850] V. missouriensis E. Greene [C 3777] There were 105 species, 71 genera, and 42 families in Andrew’s Bog. There were 118 species, 78 genera, and 42 families in Chester’s Bog. Sorensen’s Index of Similarity (Sorensen 1948) between them is 80, meaning that they are essentially identical floristically. Combining both lists, dicots account for 45% of the total. Monocots, ferns, and others account for 55% of species, which is typical of bog communities elsewhere. The vast majority of plant species at both study sites were hydrophytes (Reed 1988). The main families were Cyperaceae, Juncaceae, Poaceae, Xyridaceae, Apiaceae, and Asteraceae. Steve Orzell and Edwin Bridges made extensive observations and collections on these bogs in 1988 and 1990. They have vouchers for a number of species that we did not find. In Table 2, we list these alphabetically and indicate which bog they came from. We did not include these collections in our list because they were made a decade ago and might not presently be at these sites and because we collected only in a specific portion of each bog. Orzell & Bridges’ collection numbers are given, but their collections have yet to be deposited in an herbarium. Absence of a specimen number indicates that the taxon was not collected but only entered in field notes. Table 2. Orzell & Bridges’ additions to Chester’s (C) and Andrew’s (A) bog flora. Numbers refer to Bridges & Orzell collections. Aletris aurea L. [A] Amorpha paniculata Torrey & Gray [A 7961] Carex longii Mack. [A 13694, C 13721] Cirsium muticum Michx. [A 7974] Coreopsis tripteris L. [A 7970] Eleocharis quadrangulata (Michx.) Roem. & Schult. [A 7958] Ludwigia linearis Walt. [A 7967] L. sphaerocarpa Ell. [A 7956] Melanthium virginicum L. [A] Mitreola sessilifolia (Gmel.) G. Don [A 7952] Panicum hemitomon Schult. [A] Rhynchospora caduca Ell. [A 7971] R. stenophylla Chapm. ex Curtis [A 7184] Schizachyrium scoparium [A 7959] Scleria triglomerata Michx. [C 13729] Sorghastrum nutans (L.) Nash [A] Thelypteris palustris Schott [A 6590] MacRoberts & MacRoberts: Floristics of muck bogs 69 Xyris laxifolia Matt. var. iridifolia (Chapm.) Kral [A] Table 3 gives information on five soil samples from Chester’s and Andrew’s bogs. The “sand” samples were taken near the bog edge or margin, “med” was taken in the bog proper, and “deep” was taken about 35 cm below the surface in the bog proper. Table 3. Soil chemistry of Chester’s and Andrew’s bogs. “Sand” samples were taken near the bog edge or margin; “med” was taken in the bog proper; and “deep” was taken about 35 cm below the surface in the bog proper. ao CCE Xchangeable Ions (ppm) J) et ee |Andrew'sSand | 46 | 7 | 41 | 200 | 4 JT 3. |Andrew'sDeep | 48 | 8 | 75 | 769 _| |ChestersMed | 43 | 8 | 18 | 12 [Chester's Deep {| 48 | 13) [ 69 | 115 A more detailed analysis conducted on the Chester’s Deep sample revealed the following dry basis: N = 1.71%, P = 0.035%, K = 0.29%, S = 0.53%, Mg = 0.21%, Ca = 0.59%, Na = 361 ppm, Fe = 5130 ppm, Al = 43200 ppm, Mn = 50 ppm, Cu < 0.01 ppm, Zn = 75 ppm. Organic matter, measured by combustion, was 50.4%. (Apparently the calorimetric method is accurate only for low organic content samples; consequently both Andrew’s and Chester’s bogs undoubtedly have high Organic matter in the deep, muck/slurry areas.) DISCUSSION Rowell (1949a, 1949b), Kral (1955), and Starbuck (1984) have developed checklists for muck bogs in the post oak savanna region. Rowell’s study site was in Robertson County about 115 km SSW of our study site, and Kral’s sites were in Van Zandt and Freestone counties about 57 km NNE and S50 km SSW, respectively, of our study sites. Starbuck (1984) studied several bogs in Robertson County about 100 km SSW of Anderson County. To make comparisons with our study sites, we modified some of these lists to exclude species that were obviously members of other associations. Kral lists 80 species for his “Bog Edge” and “Bog Proper”. Of these, 52 or 65% occurred in the bogs we studied. Rowell’s (1949a) list is somewhat more problematic, but we found that of 68 species from his “zone two,” which consisted of “low shrubs, grasses, and other herbaceous forms”, 45 or 66% occurred at the two 70 PHYTOLOGIA July 1998 volume 85(1):61-73 Anderson County bogs. Starbuck (1984) lists 78 “bog species” of which we found 46 or 59% in Chester’s and Andrew’s bogs. Although these figures should be considered tentative since sampling methods among these studies differed, they nonetheless point to strong affinities among these different sites. On the other hand, muck bogs show marked differences to pitcher plant bogs in deep east Texas and west Louisiana (Nixon & Ward 1986; Orzell 1990; MacRoberts & MacRoberts 1988, 1993), with a Sorensen’s Index of Similarity of between 43 and 48. Thus, while both hillside bogs and post oak savanna muck bogs have many species in common, for example, Sarracenia alata and Eriocaulon decangulare, many — others are not shared. The post oak region is the western range limit of many bog species (e.g., Sarracenia alata, Lycopodiella appressa, Burmannia capilata, and Eriocaulon decangulare), and many other species drop out of the piney woods flora long before the post oak savanna region is reached (e.g., Magnolia virginiana, Persea borbonia, Pinguicula pumila, and Rhexia lutea). Also, because of habitat conditions, in the post oak savanna region, marsh species intermingle with bog species to create a unique association. Classifications of Texas vegetation by region and by community fail to capture the nature of the muck bog plant associations or, for that matter, any of the inland marsh communities. Of all the wetland communities described by Correll & Correll (1972), inland marshes and bogs appear to be the most understudied. While the report “Preserving Texas’ Natural Heritage” (LBJ 1978) describes “peat bogs and marshes” as distributed through the post oak savanna region, no further description is given and no information is provided about the relation between these two communities. Diamond et al. (1987) describe bogs as occurring in both the piney woods and the post oak savanna, but barely mention fresh water marshes except to say that they occur in “all regions” of Texas. The community list of the Texas Natural Heritage Program (1992) does no better; it barely mentions inland marshes, only indicating that Juncus is a dominant genus and that marshes occur throughout Texas. The Nature Conservancy classification of ecological communities of the southeastern United States (Weakley et al. 1998) does not describe Texas muck bogs: although many marsh communities that resemble Texas post oak savanna marshes are listed, no Texas sites are specifically mentioned. Although our experience of wetland communities in the post oak savanna region of the WGCP is limited, at present we believe that, while there were few natural lakes, historically there were many marshes, hillside bogs, muck bogs, peatlands, and probably other kinds of marsh-bog herb-dominated wetlands. What we have described for the GEWMA sites appears to be an interdigitation or intermingling of hillside bog and marsh. This appears to be what Rowell (1949a), Kral (1955), and Lodwick (1975) have described. This intermingling would explain the very high biomass found by Lodwick (1975) when compared with bogs elsewhere (Allen et al. 1988). In conclusion, bogs and marshes and the bog-marsh complex of the post oak savanna region of Texas need additional and detailed research and sampling before any understanding of the wetlands in this area will be gained. Especially important are MacRoberts & MacRoberts: Flonstics of muck bogs 71 floristic studies of those hillside bogs that do not grade into marshes, and studies of inland marshes in general. ACKNOWLEDGMENTS Jim Cathey, Donnie Frels, and Hayden Haucke of the Gus Engeling Wildlife Management Area aided with the study. Susan L. Grace, National Wetland Research Center, Lafayette, Louisiana aided in various ways. Carl Frentress, Texas Parks and Wildlife, showed us sites in Henderson County. Financial support was provided in part by a USGS Biological Resources Division species at risk grant to the National Wetlands Research Center. Larry Brown helped with plant identifications, especially grasses and Fuirena. Guy Nesom aided with Asteraceae. Steve Orzell shared with us his and E.L. Bridges’ data on Andrew’s and Chester’s bogs. Jim Cathey, Sue Grace, Robert Kral, and Steve Orzell read an earlier version of this paper. LITERATURE CITED Allen, C.M. C.H. Stagg, & S.D. Parris. 1988. Analysis of the vegetation in pitcher plant bogs in two baygalls at Fort Polk in west central Louisiana. Proc. Louisiana Acad. Sci. 50:1-6. Bridges, E.L. & S.L. Orzell. 1989a. Longleaf pine communities of the west gulf coastal plain. Natural Areas Journal 9:246-263. Bridges, E.L. & S.L. Orzell. 1989b. Additions and noteworthy vascular plant collections from Texas and Louisiana, with historical, ecological, and geographical notes. Phytologia 66: 12-69. Bryant, V.M. 1977. A 16,000 year pollen record of vegetational change in central Texas. Palynology 1:143-156. Correll, D.S. & H.B. Correll. 1972. Aquatic and Wetland Plants of Southwestern United States. Stanford University Press, Stanford, California. Diamond, D.D., D.H. Riskind, & S.L. Orzell. 1987. A framework for plant community classification and conservation in Texas. Texas J. Sci. 39:203-221. Folkerts, G.W. 1991. A preliminary classification of pitcher plant habitats in the southeastern United States. J. Alabama Acad. Sci. 62:199-225. Graham, A. & C. Heimsch. 1960. Pollen studies of some Texas peat deposits. Ecology 41:751-763. Harcombe, P.A., J.S. Glitzenstein, R.G. Knox, S.L. Orzell, & ELL. Bridges. 1993. Vegetation of the longleaf pine region of the west gulf coastal plain. Proc. Tall Timbers Fire Ecology Conf. 18:83-104. Haucke, H. & J.H. Rose. n.d. Wildlife Management: Past, Present, and Future. A field guide to demonstrations of wildlife management and practices and principles on the Engeling Wildlife Management Area. Texas Parks and Wildlife Department, Austin, Texas. Johnson, F.L. 1986. Oak-hickory savannas and transition zones: Preservation status and management problems. Pp. 345-347. In: D.L. Kulhavy & R.W. Conner (eds.). Wilderness and Natural Areas in the Eastern United States: A Management Ve PHYTOLOGIA July 1998 volume 85( 1):61-73 Challenge. Center for Applied Studies, School of Forestry, Stephen F. Austin State University, Nacogdoches, Texas. Jones, S.D., J.K. Wipff, & P.M. Montgomery. 1997. Vascular Plants of Texas: A Comprehensive Checklist Including Synonymy, Bibliography, and Index. University of Texas Press, Austin, Texas. Kartesz, J.T. 1994. A Synonymized Checklist of the Vascular Flora of the United States, Canada, and Greenland. Timber Press, Portland, Oregon. Kral, R. 1955. A floristic comparison of two hillside bog localities in northeast Texas. Field & Lab. 23:47-69. Larson, D.A., V.M. Bryant, & T.S. Patty. 1972. Pollen analysis of a central Texas bog. Amer. Midl. Naturalist 88:358-367. LBJ School of Public Affairs. 1978. Preserving Texas’ natural heritage. LBJ School Publ. Aff. Pol. Res. Proj. Rept. 31:17-34. Lodwick, L.N. 1975. Net aerial primary production of three east Texas bogs. M.S. thesis, Baylor University, Waco, Texas. MacRoberts, B.R...& M.H. MacRoberts. 1988. Florstic composition of two west Louisiana pitcher plant bogs. Phytologia 65: 184-190. MacRoberts, B.R. & M.H. MacRoberts. 1990. Vascular flora of two west Louisiana pitcher plant bogs. Phytologia 68:271-275. MacRoberts, B.R...& M.H. MacRoberts. 1991. Floristics of three bogs in western Louisiana. Phytologia. 70:135-141. MacRoberts, B.R. & M.H. MacRoberts. 1992. Floristics of four small bogs in western Louisiana with observations on species-area relationships. Phytologia 73:49-56. MacRoberts, B.R. & M.H. MacRoberts. 1993. Floristics of a bog in Vernon Parish, Louisiana, with comments on noteworthy bog plants in westem Louisiana. Phytologia 75:247-258. Nesom, G.L. 1994. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the new world species. Phytologia 77:141-297. Nixon, E.S. & J.R. Ward. 1986. Floristic composition and management of east Texas pitcher plant bogs. Pp. 283-287. In: D.L. Kulhavy & R.W. Conner (eds.). Wilderness and Natural Areas in the Eastern United States: A Management Challenge. Center for Applied Studies, School of Forestry, Stephen F. Austin State University, Nacogdoches, Texas. Potzger, J.E. & B.C. Tharp. 1943. Pollen record of Canadian spruce and fir from Texas bogs. Science 98:584. Potzger, J.E. & B.C. Tharp. 1947. Pollen profile from a Texas bog. Ecology 28:274-279. Potzger, J.E. & B.C. Tharp. 1954. Pollen study of two bogs in Texas. Ecology 35:462-466. Reed, P.B. 1988. National list of plant species that occur in wetlands: National summary. U.S. Fish and Wildlife Service, Washington, D.C. Rowell, C.M. 1949a. Floral composition of a Sphagnum bog in Robertson County, Texas. M.S. thesis, Texas A&M University, College Station, Texas. Rowell, C.M. 1949b. A preliminary report on the floral composition of a Sphagnum bog in Robertson County, Texas. Texas J. Sci. 1:50-53. Sorensen, T. 1948. A method of establishing groups of equal amplitude in plant sociology based on similarity of species content. Pp. 234-249. In: R.P. McIntosh (ed.). Phytosociology. Benchmark Papers in Ecology. Dowden, Hutchinsom & Ross, Stroudsburg, Pennsylvania. MacRoberts & MacRoberts: Floristics of muck bogs i e Starbuck, T.J. 1984. The vascular flora of Robertson County, Texas. M.A. thesis, Texas A&M University, College Station, Texas. Telfair, R.C. 1988. Conservation of the Catfish Creek ecosystem: A national natural landmark in eastern Texas. Texas J. Sci. 40:11-23. Telfair, R.C., J.H. Rose, & G.H. Veteto. n.d. Vegetation of the Gus Engeling Wildlife Management Area. Texas Parks and Wildlife Department, Austin, Texas. Texas Natural Hentage Program (TNHP). 1992. Plant communities of Texas (Series Level). Unpublished report, Texas Parks and Wildlife Department, Austin, Texas. Weakley, A.S., K.D. Patterson, S. Landall, M. Pyne, and others (compilers). 1998. International classification of ecological communities: Terrestrial vegetation of the southeastern United States. Working draft. The Nature Conservancy, Chapel Hill, North Carolina. Phytologia (July 1998) 8(1):74-79. SEX MORPH DESCRIPTIONS OF MALOSMA LAURINA (ANACARDIACEAE), A POLYGAMODIOECIOUS SPECIES Gary B. Perlmutter 151 North Lomita Avenue, Ojai, California 93023 U.S.A. ABSTRACT In 1994-1998 I examined laurel sumac (Malosma laurina) shrubs in Matilija Canyon, Ventura County, California for sex-identifying characteristics, augmented by study of herbarium specimens. Malosma is confirmed as polygamodioecious with detailed descriptions of staminate, andromonoecious and pistillate sex morphs at the individual, inflorescence, flower and organ levels. Characteristics are presented for identifying sex morphs of individuals in the field. KEY WORDS: Malosma, Anacardiaceae, polygamodioecy INTRODUCTION Malosma laurina (Nutt. ex T. & G.) Nutt. ex Abrams (laurel sumac) is a large, evergreen shrub found commonly in coastal sage scrub and chaparral in southern and Baja California. Although it has been described as polygamous or polygamodioecious since its first published treatment (Torrey & Gray 1838; Brewer & Watson 1876; Abrams 1911; Barkley 1937; Wilken 1993), descriptions of unisexual and bisexual flowers or other sex-identifying characteristics of individual plants are largely lacking. The only distinctive description is of pistillate plants, from Engler (1883). The purpose of this study is to provide detailed descriptions of flower and inflorescence morphology, to verify the breeding system of M. laurina as polypamadion=s and provide a tool for identifying sexes in the field. 3 STUDY SITE Field studies were conducted in Matilija Canyon, just west of the junction of Matilija Creek and its Upper North Fork, in the eastern end of the Santa Ynez 74 Perimutter. Sex morphs in Malosmalaurina ge Mountains at 490-670 m, 29 km NNW of Ventura, California (34° 03’ N, 119° 22’ W). Located 17 km from the coast, this area is in the Transition Climate Zone (Bailey 1966), influenced by a mixture of maritime and continental air masses (Hickman 1993). At MatilijaDam, 7 km ESE, mean annual rainfall was 91.4 cm for 1950-1985. Mean January minimum and August maximum temperatures were 3.8 and 33.3°C, respectively. Topographically, the area includes the canyon floor and a south-facing slope. The vegetation is a mosaic of chaparral dominated by Adenostoma fasciculatum, Ceanothus crassifolius, and M. laurina, ranging in height from 1-3 meters. Fire history includes burns in 1985 and 1993. METHODS While tracking inflorescences (or “thyrsi,” after Barkley [1937]) of twenty study plants for a phenology study in 1994, I discovered differences in flowers and differential fruiting both in timing and production among individuals. This allowed the classification of individual shrubs as “staminate,” “pistillate,” and “andromonoecious” (the lattermost following Richards [1986]). Three voucher specimens representing shrubs of each above sex (G.B. Perlmutter s.n. [1995] each) were deposited at SBBG. In 1995, I measured lengths and widths of twenty thyrsi (five replicates from four staminate and four pistillate plants each) in the field and analyzed them using the Student’s t test to determine significant differences between the two sets. In 1997, flowers were examined on plants of each of the three sexes, noting qualitative morphological characteristics. For quantitative analysis at the organ level, anthers of four staminate and two pistillate herbarium specimens at SBBG (n, = 10) were measured. The resulting six data sets were then tested for differences by single factor ANOVA (Model I) followed by Tukey multiple companson tests (Zar 1984). Staminate specimens included Hoffman s.n. (1927), Philbrick s.n. (1972-1973), Pollard s.n. (1969), and Pollard (1968); pistillate ones were Murphy & Newman s.n. (1981) and Ackley s.n. (1928). Level of significance for all statistical tests were set a priori at 0.05. In 1998, fertilization tests were conducted on one andromonoecious and one pistillate shrub. On each plant one thyrsus was left unbagged and untreated to serve as a control; experimentally manipulated inflorescences were bagged to prevent natural pollination by insects. On the andromonoecious shrub two bagged inflorescences were tested for self-pollination (flowers were artificially pollinated by lightly brushing the inflorescence with a small paint brush, using pollen from the same flowers), and two others on this same plant were bagged only to test for self-fertilization by autogamy. Two thyrsi on the pistillate plant were similarly bagged only to test for apomixis. Inflorescences were checked weekly for health and reproductive development. Any thyrsi showing disease were terminated early. Otherwise, tests were ended when fruits matured, at which time they were collected to determine fruiting success by dividing their fruit number by an estimated bud number for inflorescences of similar size. Bud number estimates were developed from a linear regression equation (y = 49.221x - 518.31) relating panicle size (x = length x width, 76 PHYTOLOGIA July 1998 volume 85(1):74-79 in cm?) with bud number (y), from a sample of 20 inflorescences with size and corresponding bud number ranges of (22.5 - 102 cm’) and (804 - 5,698 buds), respectively. The regression equation was from a highly significant correlation of the two above variables (r° = 0.88, p < 0.01). RESULTS Individual level. Sexes were distinguishable at flowering and fruiting. Pistillate shrubs had pinkish inflorescences in flower; producing full infructescences upon one week after flowering. Staminate plants were observed to have yellowish thyrsi, which fade to tan one to two weeks after anthesis. Andromonoecious individuals appeared indistinguishable from staminate ones upon flowering, but later produced an estimated 0-25% fruit per individual (two plants had only one fruit observed) 3 weeks later than pistillate shrubs. (Fruiting in these plants varied among years, some not producing fruits in certain years [pers. obs.]). Inflorescence level. Pistillate shrubs had inflorescences with mean (+ SE) lengths of 8.4 (+ 0.5) cm (range 4.0-12.0 cm, N = 20) and mean (+ SE) widths of 4.8 (+ 0.3) cm (range 2.0-7.0 cm, N = 20). Mean (+ SE) thyrsus lengths and widths of staminate and/or andromonoecious (i.e., male-fertile) plants were 10.9 (+ 0.6) cm (range 7.5- 13.5, N = 20) and 7.2 (+ 0.5) cm (range 4.5-14.0 cm, N = 20), respectively. Both lengths and widths were larger among male-fertiles (one-tailed Student’s t tests, p < 0.005 and p < 0.001, respectively). Flower level. Three flower types were identified: pistillate, staminate, and bisexual. In pistillate flowers pistils were reddish, with styles extending above the level of the corolla. Anthers lacked pollen, appeared pale yellow and smooth, and were borne on filaments which did not extend beyond the corolla. Pistillate flowers were uniform throughout the plant. . Staminate flowers had shorter pistils, which lacked color and were often obscured by clustered stamens. Anthers produced pollen, appeared bnght yellow, and the filaments extended well beyond the corolla. Staminate flowers often dominated the inflorescence, either occurring exclusively or mixed with bisexual flowers. Bisexual flowers had reddish pistils like pistillate flowers. Stamens radiated from each carpel, having anthers which produced pollen. Fruits developed three weeks later than those on pistillate shrubs. Bisexual flowers occurred variably (up to 25%) in inflorescences with staminate flowers, sometimes singly on a whole plant (pers. obs.). Likewise, one to nearly all thyrsi with mixed flowers were observed on an apparently staminate plant. Self-fertilization tests resulted in only the pistillate control inflorescence producing fruit: 16 drupes out of an estimated 1,586 buds or 1% fruit success. This is representative of the overall fruit success of the shrub (pers. obs.). The andromonoecious control inflorescence was likewise representative of that plant, with near complete -failure of fruit development. Therefore, the pistillate results Perlmutter. Sex morphs in Malosmalaurina 77 demonstrated non-apomixis and the andromonoecious results implied self- incompatibility. Organ level. Anther lengths ranged 0.6-1.5 mm overall, ranging 0.6-0.8 mm among pistillate flowers and 1.0-1.5 mm among staminate flowers. The ANOVA test revealed differences among specimens (among ms = 1.02, among df = 5, p < 0.005, N = 60), which were consistent between sexes (Tables 1 and 2). DISCUSSION The breeding system of Malosma laurina is confirmed to be polygamodioecious, as described by Barkley (1937). Polygamodioecy is defined as functionally dioecious plants having few hermaphroditic flowers on otherwise staminate or pistillate individuals (D. Young, pers. comm.). In this case, perfect flowers occurred only on predominately staminate shrubs. Male-sterility and female-sterility in M. laurina is based on the observation of underdeveloped organs of the opposite sex, rendering them sterile by failing to mature and function reproductively. Previous to this study, the only published account of stunted organs in M. laurina is by Engler (1883), which my findings confirm and elaborate upon. This functional sterility in morphologically hermaphrodite flowers is also found in other members of the Anacardiaceae such as Toxicodendron diversilobum (= Rhus diversiloba) in Jepson (1925) and Rhus integrifolia and R. ovata (Young 1972). It is possible that the breeding system in M. laurina is digenic, as postulated by Charlesworth & Charlesworth (1978). Further study is needed. Using the descriptions here provided, Malosma individuals can be readily sexed in the field. Pistillate shrubs can be easily identified by the pinkish color of inflorescences in flower and later by heavy production of fruit. To discern an andromonoecious plant from a staminate one, close examination of thyrsi is necessary to locate bisexual flowers or infrequent fruit. The presence of few fruits is easier to identify a plant as andromonoecious, but this characteristic should not be used alone, as an absence of fruit may be a result of pollination failure and not necessanly indicative of complete female-sterility. Therefore, it is recommended that a combination of flower and fruiting observations be used in sexing a male-fertile shrub as either staminate or andromonoecious. ACKNOWLEDGMENTS Dieter H. Wilken of the Santa Barbara Botanic Garden (SBBG) was instrumental in the formation and development of this study, providing advice and direction throughout. SBBG Librarian Laurie Hannah was helpful by providing me with early published treatments of Malosma. David A. Young of Colorado State University along with D. Wilken offered many instructive comments on earlier versions of this manuscript. [ also thank Richard Colla of the MatilijaCanyon Ranch for allowing me 78 PHYTOLOGIA July 1998 volume 85(1):74-79 Table 1. Means and standard errors of Malosma laurina anther lengths from six ranked SBBG specimens (n; = ‘0). Ranking was for Tukey multiple comparison tests (Table 2). LO Sn ae ES, Po ee H.M. Pollard s.n. (1968) 1.1 (40.04) R.N. Philbrick s.n. (1972-1973) 1.2 (40.03) R. Hoffman s.n. (1927 1.3 (40.05 H.M. Pollard s.n. (1969 1.4 (40.02) T. Murphey / J. Newman s.n. (1981 0.7 (40.02 M.N. Ackley s.n. (1928) 0.7 (£0.04) Table 2. Tukey multiple comparison tests of Malosma laurina anther lengths from six SBBG specimens (n; = 10). Comparison p value Ea ee ae < 0.001 < 0.001 < 0.001 < 0.05 < 0.001 < 0.001 < 0.001 _<0.001 < 0.001 < 0.001 < 0.001 Perlmutter. Sex morphs in Malosma laurina 79 to study on his property, the Matilija Canyon Ranch - Wildlife Sanctuary. Steven G. Lawry of Lawry's Technical Services provided transportation and support. Climatic data were provided by the Ventura County Flood Control District. LITERATURECITED Abrams, L.R. 1911. Flora of Los Angeles and Vicinity (supplemented ed.). Stanford University Press, Stanford, California. Bailey, H.P. 1966. Weather of Southern California. University of California Press, Berkeley, California. Barkley, F.A. 1937. “A monographic study of Rhus and its immediate allies in North and Central America, including the West Indies.” Ann. Missouri Bot. Garden 24:265-498 Brewer, W.H. & S. Watson. 1876. “Polypetalae.” Botany of California, Volume 1. Welch, Bigelow, and Co., University Press, Cambridge, Massachusetts. Pp. 1- 276. Charlesworth, B. & Charlesworth, D. 1978. A model for the evolution of dioecy and gynodioecy. Amer. Naturalist 112:975-997. Engler, A. 1883. “Burseraceae et Anacardiaceae.” Jn: Candolle, A.L.P.P. de & A.C.P. Candolle. Monographiae Phanerogamarum Prodromi nunc Continuatio, nunc Revisio. 4:393. Jepson, W.L. 1925. A Manualof the Flowering Plants of California. University of California Press, Berkeley, California. Hickman, J.C. 1993. The Jepson Manual: Higher Plants of California. University of California Press, Berkeley, California. Richards, A.J. 1986. Plant Breeding Systems. George Allen and Unwin, London, United Kingdom. Torrey, J. & A. Gray. 1838. A Flora of North America. Hafner Publishing Company, New York, New York. Wilken, D.H. 1993. “Anacardiaceae.” In: J.C. Hickman. The Jepson Manual: Higher Plants of California. University of California Press, Berkeley, California. Pp. 134-136. Young, D.A. 1972. The reproductive biology of Rhus integrifolia and Rhus ovata (Anacardiaceae). Evolution 26:406-14. Zar, J.H. 1984. Biostatistical Analysis (2nd ed.]. Princeton-Hall, Englewood Cliffs, New Jersey. Phytologia (July 1998) 85(1):80. BOOKS RECEIVED Physiological Plant Ecology. Third edition. Walter Larcher. Springer-Verlag New York, Inc., 160 Imlay St., Brooklyn, New York 11231. 1995. xvi. 506 pp. $44.50 ISBN 0-387-581 16-23 (hardcover). After an introductory chapter on the environment in which plants grow, this book has extensive chapters on carbon metabolism, other nutrient metabolism, water relations, growth and development (as influenced by environment), and effects of various stresses on plants. Phytochemistry of Plants Used in Traditional Medicine. K. Hostettmann, A. Marston, M. Maillard, & M. Hamburger (eds.). Proceedings of the Phytochemical Society of Europe, vol. 37. Oxford Science Publications, Clarendon Press, Oxford, Oxford University Press, 198 Madison Avenue, New York, New York 10016. 1995. xiv. 408 pp. $130.00 ISBN 0-19-857775-3 (hardcover). A compendium of sixteen articles composed by 34 authors is presented in this volume. The authors and their works contained within this book represent efforts to understand traditional medicinal practices from a large diversity of cultures around the world. These papers are based on presentations made at a symposium in October 1993. 80 Phytologia (July 1998) 85(1):81. PUBLICATION DATES, VOLUME 84 Issue Cover date Publication date 84(1) January 1998 16 December 1998 84(2) February 1998 26 February 1999 84(3) March 1998 14 June 1999 84(4) Apnil 1998 29 July 1999 84(5) May 1998 26 August 1999 84(6) June 1998 27 September 1999 81 Phytologia (July 1998) 85(1):82. INDEX TO REVIEWERS, VOLUME 84 The editor expresses his most sincere appreciation to the following individuals. These are persons who have reviewed papers that were submitted for publication in volume 84 of Phytologia. Without the willingness and diligence of these reviewers, the task of the editor would be much more difficult, and the quality of the papers published would be lessened. To each of you, I offer my most sincere thanks. Michael J. Warnock, Editor Brown, | 2 Nesom, G.L. Croat, T.B. Nicolson, D.H. Delevoryas, T. Pruski, J.F. Demeke, T. Rosenzweig, N. Duell, E.A. Scharpf, R.F. Eggli, U. Singhurst, J. Endress, M. Sivyer, S.I. Evans, R. Teague, J. Filip, G.M. Thomas, R.D. Gandhi, K.N. Turner, B.L. Gorst van Rijn, A.R.A. Turner, G. Grant, K.A. Wendt, T. Hammel, B.E. Wiersema, J. Judd, W. Wilken, D. Kral, R. Williams, J. Lellinger, D.B. Young, D.A. Liggio, J. MacRoberts, B.R. MacRoberts, M.H. Massey, J. McDaniel, S. Phytologia (July 1998) 85(1):83. INDEX TO AUTHORS, VOLUME 84 Adams, R.P. 354 ter Welle, B.J.H. 354 Allorge-Boiteau, L 304 White, H.L. 93 Beatty, J.S. 154 Branch, J.R. 93 Brown, L.E. 107 Day, A.G. 368 Doweld, A.B. , 363 Geils, B.W. 154 Grant, V. 69, 368 Grayum, M.H. 307 Hershkovitz, M.A. 98 Holmes, W.C. 93 Isikov, V.P. 328 Laferniére,J.E. 169 MacRoberts, B.R. 1, 38, 297 MacRoberts, M.H. 1, 38, 297 Mathiasen, R.L. 154 Nesom, G.L. 43, 50, 107 Parks, C.G. 154 Reveal, J.L. 363 Robinson, H. 40, 345, 347 Rosenzweig, N. 28 Turner, B.L. 61, 87 Volz, P.A. 28, 328 Walker, S.B. 64 Ward, D.B. 385 Warnock, M.J. 172 83 Phytologia (July 1998) 85(1):84. NEW NAMES IN THIS ISSUE OF PHYTOLOGIA As a result of the International Botanical Congress in Tokyo in 1993, the International Association of Plant Taxonomy has been tasked with exploring the feasibility of registration of plant and fungi names. In accordance with terms of the pilot implementation of the registration concept, new names and combinations produced in this issue of PHYTOLOGIA are listed below. New name or combination Page Number Agnorhiza (Jepson) W.A. Weber, gen. nov. 19 Agnorhiza bolanderi(A. Gray) W.A. Weber, comb. nov. 19 Agnorhiza elata (H.M. Hall) W.A. Weber, comb. nov. 19 Agnorhiza invenusta (Greene) W.A. Weber, comb. nov. 20 Agnorhiza ovata (Torrey & Gray) W.A. Weber, comb. nov. 20 Agnorhiza reticulata (Greene) W.A. Weber, comb. nov. 20 Balsamorhiza lanata (W.M. Sharp) W.A. Weber, comb. nov. 20 Helianthella californica A. Gray subsp. nevadensis (Greene) W.A. Weber, comb. nov. 20 Helianthella californica A. Gray subsp. shastensis (W.A. Weber) W.A. Weber, comb. nov. 20 Scabrethia W.A. Weber, gen. nov. 20 Scabrethia scabra(W.J. Hooker) W.A. Weber, comb. nov. 21 Scabrethia scabra (W.J. Hooker) W.A. Weber subsp. attenuata (W.A. oa W.A. Weber, comb. nov. Scabrethia scabra (W.J. Hooker) W.A. Weber subsp. cinerea (W.A. ay W.A. Weber, comb. nov. 21 Cn 0 grate pees coe Articles from “beranieal ‘systematics. Sat ‘ecology, | jaeding | Pa biographical sketches, critical reviews, and summaries of literature — will be considered for publication in PHYTOLOGIA. Manuscripts. may @ be submitted either on computer diskette, or as clean ‘typescript. -- Diskettes will be returned to authors after action has been taken on 3 the manuscript. Diskettes may be 5.25” or 3.5” and may be written in — any IBM or Macintosh compatible format. Typescript: manuscripts. a should. be single spaced and will be read into the computer using a scanner. The scanner will read standard type. fonts. but will not read. é dot matrix print. Manuscripts submitted in dot matrix print” cannot — 3 be accepted. Use underscore (not italics) for scientific names. ~ Language of manuscripts may be either English or Spanish. Fes will be reduced to fit within limits of text pages. 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