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The Systematic Significance of Seed Morphology in Portulaca (Portulacaceae) Under Scanning Electron Microscopy Author(S): James F

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The Systematic Significance of Seed Morphology in Portulaca (Portulacaceae) Under Scanning Electron Microscopy Author(S): James F The Systematic Significance of Seed Morphology in Portulaca (Portulacaceae) Under Scanning Electron Microscopy Author(s): James F. Matthews and Patricia A. Levins Source: Systematic Botany, Vol. 11, No. 2 (Apr. - Jun., 1986), pp. 302-308 Published by: American Society of Plant Taxonomists Stable URL: http://www.jstor.org/stable/2419120 . Accessed: 09/05/2014 05:55 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Society of Plant Taxonomists is collaborating with JSTOR to digitize, preserve and extend access to Systematic Botany. http://www.jstor.org This content downloaded from 83.204.224.234 on Fri, 9 May 2014 05:55:35 AM All use subject to JSTOR Terms and Conditions SystematicBotany (1986), 11(2): pp. 302-308 ? Copyright1986 by the American Society of Plant Taxonomists The SystematicSignificance of Seed Morphology in Portulaca (Portulacaceae) under Scanning Electron Microscopy JAMES F. MATTHEWS and PATRICIA A. LEVINS Department of Biology, University of North Carolina, Charlotte, North Carolina 28223 ABSTRACT. Seeds of seven southeasternand one southwesternUnited States species of Portulaca were examined with the scanning electron microscope.Some species could be characterizedby the external seed morphology while others could not. Those species with the largest geographical ranges and ecological diversities showed the greatestvariation in morphology. Comparisons be- tween species oftenshowed significantoverlap of morphological characteristics.Seed morphology in Portulacacan be used as a taxonomic trait,but should be combined with other traits for the diagnostic determinationof species. Since Davis and Heywood (1963) proposed 2. A distinctiveseed coat morphology usually seed morphology as a potential taxonomic allowing identificationof species solely on character,an increasingnumber of papers have the basis of seed characters(Clark and Jern- reportedon the scanning electron microscopy stedt 1978). of seed surface patterns.Brisson and Peterson 3. Both inter-and intrapopulationalvariation; (1976, 1977) provide a complete bibliography some species monomorphicand some poly- through 1977. morphic with small populations tending to In general, the findingsuntil now have sup- be monomorphicand widespread taxa poly- ported the following groupingsregarding seed morphic (Hauptli et al. 1978). surfacetaxonomic implications. 4. Seed morphologybeing subject to the same uncertaintiesand ambiguities as any other 1. Intrageneric consistency complete, so that morphological attributeused in taxonomic seed surfacecharacters are predictable from studies (Newell and Hymowitz 1978). species to species (Chuan and Heckard 1972; 5. Monomorphic seeds with variation in those Hill 1976; Clark and Jernstedt1978; Crow species with the largest distribution(Canne 1979). 1979). 2. Intragenericconsistency high, some species 6. External seed morphology distinctive and variability (Newell and Hymowitz 1978; valuable for species distinctions (Wofford Canne 1979, 1980; Wofford1981; Elisens and 1981). Tomb 1983; Chance and Bacon 1984). 7. High levels of infraspecificvariation sug- 3. Intragenericvariability of some species, re- gesting caution in using SEM-level features lated to ecological conditions (Hauptli et al. to characterizespecies or taxa (Wyatt 1984). 1978). 8. Species specificseed coat morphologies that 4. Intraspecificvariability, showing different do not correspond well with the more clas- expressions over geographical range and sical treatmentsand suggest a restructuring ecological conditions (Wyatt 1984; Mat- of the intragenericand even the generic tax- thews and Levins 1985a). onomy (Chance and Bacon 1984). 5. Intraspecificconsistency, involving subspe- 9. Variability in seed surface patterns occurs cies, related to ploidy level (Danin et al. within geographic regions and within the 1978). species (Matthews and Levins 1985a). These varied findings have resulted in the A recentstudy (Matthews and Levins 1985a) following conclusions, listed in historical se- concentratedon the variation in seed surface quence. morphology of Portulacapilosa over its south- 1. SEM photographs have not provided suffi- eastern and southwesternUnited States range. cient evidence to sustain a taxonomic inno- The present report will analyze the six other vation not held before(Brisson and Peterson species occurring in the southeastern United 1976). States,which constitutethe seven southeastern 302 This content downloaded from 83.204.224.234 on Fri, 9 May 2014 05:55:35 AM All use subject to JSTOR Terms and Conditions 1986] MATTHEWS & LEVINS: PORTULACA 303 species of Portulaca (Matthews and Levins TABLE 1. Voucher data for representativespeci- 1985b), plus one species fromthe southwest. mens of Portulacaused for SEM studies. MATERIALS AND METHODS Fig. 1. P. oleracea L. Alabama: Mobile Co., 26 Aug 1965, DeramusD732 (ALU). Seeds were examined fromdried specimens Fig. 2. P. oleraceaL. Louisiana: Rapides Parish, 19 Jul fromthe followingherbaria: ALU, ASTC, DUR, 1979, Pias 4345 (NLU). FLAS, FSU, G, GA, JSU, KNK, KY, LAF, LL, Fig. 3. P. pilosa L. Florida: Citrus Co., 13 Aug 1958, MO, NCU, NLU, NO, NY, SMS, SMU, TENN, Kral 7825 (NY). TEX, UARK, UNA, UNCC, US, USAM, USCH, Fig. 4. P. pilosa L. Arkansas: White Co., 14 Oct 1974, USF, VDB, VPI, VSC, and WILLI. Representa- Demarees.n. (GA). Fig. 5. P. pilosa L. Oklahoma: Cimarron Co., 4 Aug tive specimens from which seeds were taken 1977, Taylor25255 (NLU). for microscopy are listed in table 1. For the Fig. 6. P. pilosa L. Texas: BrewsterCo., 23 Aug 1970, electronmicroscope, seeds were sputtercoated Semple408 (MO). with gold-palladium on a Hummer V and Fig. 7. P. umbraticolaKunth. South Carolina: Lancas- viewed at 100 x with a JeolJSM-35CF scanning terCo., 25 Sep 1973, Hatleyand Sowerss.n. (UNCC). electron microscope. Fig. 8. P. rubricaulisKunth. Florida: Lee Co., 21 Jul 1954, Cooley2316 (G). RESULTS AND DISCUSSION Fig. 9. P. retusaEngelm. Texas: Williamson Co., 16 Aug 1961, Walker73 (TEX). Seeds were examined with a 10 x handlens Fig. 10. P. smalliiP. Wilson. Georgia: Hancock Co., to determinewhether the surface was smooth 22 Oct 1952, Duncan 14481 (VSC). or rough. At 25 x, tubercles could be seen on Fig. 11. P. amilisSpegaz. Georgia: Echols Co., 23 Aug the dorsal spine and the lateral surfacescould 1961, Faircloth4786 (VSC). be determined to be smooth or rough, the tu- Fig. 12. P. grandifloraHooker. South Carolina: Ker- bercles producing the rough surface.However, shaw Co., 6 Oct 1957, Radford30004 (NCU). the relation of the tubercles to the remainder of the surface cells was not easily detected. In addition, the seeds of some species were cov- ered with an iridescentcoating which oftenob- 1, 2) fromthe southeast show that the surfaces scured the surfacepatterns. To betterinterpret vary fromnearly smooth to very granulate.Tu- the primary and secondary structuresof the bercles on the dorsal spines are minute or in- seeds, they were examined at 100 x with an distinct.While Legrand (1962) questioned the SEM. use of seed surfacetexture as a characterto de- Using the classificationof Barthlott(1984) for lineate varietiesof P. oleracea,Danin et al. (1978) the microstructuralfeatures of the seed sur- used seed size and seed coat morphology to faces of Portulaca,the primarystructure of the describe nine subspecies. Cytotaxonomiccom- anticlinal walls of the isodiametric seed coat parisons of these subspecies showed that each cells is S-undulate (in earlier reports,Matthews had a fairlydistinct distribution which may in- and Levins 1985a and 1985b,the term"stellate" dicate theirspecialization to certainclimatic and was used to describe the S-undulate pattern). edaphic conditions. Although there was some The secondarystructure exhibited by the seeds subspecificvariation in seed coat morphology, is produced by channelled anticlinalwalls, with seed size and/or ploidy level were used to as- periclinal walls varying fromflat (smooth sur- sist in the final identification.In our examina- faces) to convex, to papillate, to tuberculate. tion of P. oleracea,no attemptwas made to de- Portulaca oleracea. The most widespread terminehow many of theirsubspecies could be species of Portulacain the southeast,as well as identified in the southeastern United States, in the world, is P. oleracea L., a species with only thatvariation within P. oleraceawas clear- yellow flowers,flat leaves and sparse pubes- ly evident.Our figure1 is theirP. oleraceasubsp. cence. Legrand (1962) reported that the sur- nitida,figure 2 is their subsp. granulato-stellula- faces of P. oleraceaseeds varied widely. Danin ta. et al. (1978) demonstrated a correlation be- Portulacapilosa. Portulacapilosa L., the sec- tween ploidy level and seed coat morphology ond most common species in the southeast,ex- of P. oleracea.Seed surfaces of this taxon (figs. hibits a wide variation in seed surfaceand tex- This content downloaded from 83.204.224.234 on Fri, 9 May 2014 05:55:35 AM All use subject to JSTOR Terms and Conditions 304 SYSTEMATIC BOTANY [Volume 11 4~~~~3 -A-4~~~~~~~~~ I~~~~~~~~~~~~~~- in~~I 4;- FIGS. 1-6. Scanning electron micrographsof
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