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An updated taxonomic treatment of the natural hybrids of Sarracenia L. (Sarraceniaceae) PDF

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) TechnicalRefereed Contribution An updated taxonomic treatment of the natural hybrids of Sarracenia L. (Sarraceniaceae) RayNeyland • JenniferBushnell • DepartmentofBiologyandHealth Sciences • McNeese State University • Lake Charles • LA 70609 • USA • [email protected] Wannee Tangkham • Department ofAgricultural Sciences • McNeese State University • Lake Charles • LA 70609 • USA Keywords: natural planthybrids, Sarracenia Sarracenia hybrids, Sarraceniaceae. , Abstract: Presentlythere exists confusion concerningthe legitimacy andtaxonomy ofthe named natural hybrids ofSarracenia. One reason for this confusion centers on the unsettled taxonomy ofwhat may be called the S. rubra and the S. purpurea complexes. More specifically, hybrids derived from either or both ofthese two complexes are those for which the taxonomy is most unclear. Additionally, in some cases, collection data is scant or missing. Drawn from literature, collection records and field observations, a review ofthe named natural hybrids ofSarracenia is presented. The five taxa included in the S. rubra complex are all conservatively treated in the NRCS, USDA (2014) website as subspecies of S. rubra Walter. However, Mellichamp and Case (2009) divided them into three separate species (two with subspecies). McPherson and Schnell (2011) placed S. alabamensis Case & R.B. Case and S.jonesii Wherry as subspecies ofS. rubra. Mellichamp and Case’s (2009) taxonomy is based largely onmorphological and ecological fac- tors. That is, the five members of the complex have broadly disjunct ranges that correlate with distinctivemorphology. Inthatsense, itis apracticaltaxonomyalthoughithas, asyet, nomolecular- based systematic support. The question then becomes, how to treat the various hybrids from mem- bers ofthis complex? That is, should the members ofthe S. rubra complex be treated with some level ofdistinction or should they be treated as mere variants ofa single species as much ofthe molecular evidence currently suggests? For purposes of this paper, the five members ofthe S. rubra complex will be recognized as distinct in keeping withthe taxonomy ofMellichamp and Case (2009). This decision is supported largely by practical considerations. Specifically, the recognizable morphology ofeach, correlated with their respective disjunct ranges supports their distinction based on traditional and ecological grounds. Incontrast, molecularevidence supportsthe distinctionamongthreemembersoftheS.purpurea & complex(Neyland Merchant2006, Ellisonetal. 2012).Therefore, Mellichamp andCase’s (2009) taxonomy that distinguishes S.purpurea L. subsp. venosa (Raf.) Wherry, S.purpurea subsp.pur- purea andS. roseaNaczi is well grounded. It is noted, however, thatMcPhersonand Schnell (2011 placed S. rosea as a variety ofS.purpurea subsp. venosa. It is the goal ofthispapertoprovide abriefhistory anddescriptionofthe namednaturalhybrids ofSarracenia. Forpurposes ofthispaper,this listconsists ofthe 20naturalhybrids listedbyEllison etal. (2014). 4 Carnivorous Plant Newsletter Materials and Methods This study is derived from literature and herbaria searches. Additionally, information from field observations andnew collections bythe authors augment this studyA briefdiscussion ofeachrec- ognized natural hybrid is listed in the following section. Specimens were collected using standard herbarium techniques. Taxonomy follows Mellichamp and Case (2009). Herbarium abbreviations followThiers (2014). Results andDiscussion From this study, information on 20 named natural hybrids ofSarracenia is presented. This set ofhybrids follows Ellisonetal. (2014). The following alphabetic list includestheparentnames, the A type locality and type specimen for each. briefdescription with historical and updated informa- tion is included. S. x ahlesiiC.R.Bell & F.W.Case Parents: (S. alata (Alph. Wood) Alph. Wood x S. alabamensis Case & R.B.Case subsp. wherryi (D.E.Schnell) R.B.Case. Type locality: Wet meadow, five miles southeast ofFruitdale, Washington County,AL, 1955. Holotype: Bell 1495 (NCU). Mature leaves are erect and reddish. Sepals are green and suffused with maroon; the petals are yellow with maroon shading (Bell & Case 1956). The holotype, which can be viewed online, consists ofthree flowers and no leaves. Bell and Case also tentatively identified a collection (Bell 1456a) from nearAgricola, MS as S. x ahlesii. However, that specimen was later identified as S. x exornataandcurrentlyresidesintheNCUherbarium.Noothercollectionsareknowntotheauthors. S. x areolata Macfarlane Parents: (S. leucophylla Raf. x S. alata). Lectotype: Macfarlane s.n. (NCU). Type locality: Theodore, Mobile County,AL, 1909. Erect leaves are mostly green with darkerveins and prominent white areoles on the operculum and peristome region. Sepals are mostly green and suffused with red; the petals are typically red. This hybrid is vegetatively similar to S. x moorei. Because the parents freely hybridize and their & progenyeasilybackcross (Bell Case 1956), introgressionproduces amaterial amountofbothleaf venation and flower colorvariation. Commonwithin its small range, this hybridhas been collected inJackson County, MS andMobile County,AL. The lectotype can be viewed online. S. x caseiMellichamp Parents: (S.psittacina Michx. x S. alabamensis subsp. wherryi). Type locality:Along US 45just north ofDeerPark, Washington County,AL, 1985. Holotype: Mellichamp s.n. (UNCC). The semi-decumbent leaves ofthis hybrid are thin textured and faintly hairy. Young leaves are somewhat bronze colored like those ofits S. alabamensis subsp. wherryi parent (pers. obs.). The opercula are bonnet-shaped with translucent areoles derived from its S.psittacina parent. Flowers are small (Mellichamp 2008) with (presumably) redpetals and sepals. No informationwithrespect to fragrance is known. The senior authorinBaldwin County,AL discovered a secondpopulation in Volume 44 March 2015 5 2014. Because both ofits parents bloom at about the same time in early May, this hybrid is readily produced. However, because ofthe restricted range ofS. alabamensis subsp. wherryi this hybrid is rare. Its bonnet-shaped operculum is a character also evinced in S. xformosa, S. x gilpini andS. x wrigleyana. We notethatamongeachofthesehybridswithbonnet-shapedopercula, some individu- al leaves do not completely open along the suture lines. Whenthis occurs, the leaves appearmostly closed like theirS.psittacina parent. Bothtypes ofleaves can occur onthe same individualplant. S. x catesbaeiElliott Parents: S.flava L. x S.purpurea subsp. venosa). ( Type locality: Along the margins ofrivulets amidstthehigh sandhills ofChesterfielddistrictin SC. Neotype: Macbride s.n. (CHARL). With spreading, collar-shaped opercula, the semi-decumbent leaves are marked with red or purple. Each leaftypically exhibits a dark splotch ofpurple or red on the base ofthe operculum, a characteristic ofits S.flava parent. Sepals and petals are red, a characteristic ofits S. purpurea subsp. venosa parent. However, Bell (1952) notedthatpetals mighthave tints ofyellow, acontribu- tionfromS.flava. Eliotand Stauffer(1951)provide agoodimage ofthis hybridinits nativehabitat. Althoughplantshave beencollectedoveralarge rangethatincludesthePiedmontandCoastalPlain ofGA, SC andNC, this hybridis notcommon (Bell 1952; Mellichamp & Case 2009). Its spreading operculum (not pinched in the middle), is a character also evinced in the vegetatively similar S. x chelsonii, S. x exornata, andS. x swaniana. Although the neotype collection sheet is not dated, the collectionwas made inthe early 1800s as it is referenced (as a species) by Elliot (1824). The actual specimen remains in good condition. Additional collections were located in the UNC, UNCC, and US herbaria. S. x charlesmooreiMellichamp Parents: (S.jonesiiWherry x S.purpurea subsp.purpurea). Type locality: Henderson County, NC, 2007. Holotype: Mellichamp s.n. (UNCC). Reddenedorpurplishleaves are semi-decumbentwithundulatedcollar-shapedoperculathatare pinched in the middle. Sepals and petals are red. This hybrid has been collected in three counties in the mountains ofNC. Plants collected by Wherry in Henderson County were included in Bell’s (1952) neotypification ofS. x chelsonii. However, with the acceptance of S.jonesii as a distinct species, Mellichamp (2008) recognized S. x charlesmoorei as a distinct taxon and separate from S. x chelsonii. This hybrid is rare due to the loss ofthe montane bogs inhabited by its S.jonesii parent (Mellichamp 2008). Pinched in the middle, the opercula ofthis taxon are reminiscent ofa traditional Dutch woman’s cap. This shape is a character also evinced in the vegetatively similarS. x mitchelliana andS. x naczii. S. x chelsoniiVeitch ex Masters Parents: (S. rubra subsp. rubra x S.purpurea subsp. venosa). Type locality: Eastern shore ofScotlandLake, Scotland County, NC, 1947. Neotype: Wellmans.n. (NCU). The leaves of this hybrid are semi-decumbent with spreading, collar-shaped opercula. Sepals and petals are red. Attributed to a horticulturally engineered hybrid, the name S. x chelsonii first appeared in the literature in 1878. A natural specimen was first discovered in 1933 by Wherry in southeastern North Carolina (Bell 1949). The specimen collected in 1947 by Wellman from the 6 Carnivorous Plant Newsletter western edge ofthe coastal plain in Scotland County, NC was designated as the neotype by Bell (1952). This collectionconsists ofa single leaf.An additional specimen, also collected byWellman at the type locality, consists ofa single leafand a single flower. Housed at NCU, these two are the only known specimens ofthis natural hybrid. The neotype maybe viewed online. S. x courtiiVeitch exWilson Parents: (S.purpurea x S.psittacina). Plants from this cross are known from horticulture. However, Sheridan and Scholl (1993) re- portedthat theyhad observed anatural population ofabout ten individuals ofwhat they calledS. x courtii along a road embankment in Liberty County, FL in 1989. Because such a hybrid observed here would have S. rosea as its parent (not S. purpurea), these plants would be distinct from S. x courtii. Because no collection material apparently was made, this hybrid reported from Florida remains unconfirmed. The typically large temporal difference in the blooming times ofthe parents presumablywouldmake such a hybrid, ifit exists, quite rare. S. x excellensW.Bull. Parents: (S. leucophylla x S. minor). Type: None. This hybrid was first named and described by Nicholson (1887). Commonly produced in hor- ticulture, plants are erect with prominent areoles and somewhat arching opercula. The natural oc- currence ofthis hybrid has been reported from the small area in the Florida Panhandle where the ranges ofthe two respective parentsjust barely overlap. However, our efforts to locate any natural collectionmaterial were unsuccessful. S. x exornata W.Bull. Parents: (S. alata x S. rosea). Type locality: Theodore, Mobile County,AL, (yearunknown). Neotype: Pennell s.n. (PENN). Prominently marked with red or purple veins, each leaf is semi-decumbent with a spreading, collar-shaped operculum. Bell (1952) describedthe flowers as havingred-brown sepals (sometimes tinged with green) and petals that are either red or rose with yellow margins. This hybrid has been collected inMobile County,AL nearTheodore and Spring Hill (Bell 1952). Bell also collectedthis hybridinabogfivemiles southofAgricola, MS inGeorge County(Bell & Case 1956). The USDA, NRCS (2014) distribution map indicates that this hybrid also occurs in Washington County, MS. In 2014, the senior author observed a population in Jackson County, MS. Several specimens are housed at the NCU herbarium. The neotype was located in the PH herbarium. It is noted that the PENNherbariumwas incorporated into the PH herbarium in 1974. S. xfarnhamii Farnham nothosubsp. bellii (Mellichamp) Neyland, Bushnell & Tangkham comb, et stat. nov. Basionym: S. x belliiMellichamp, Carniv. PI. Newslett. 37: 114 (2008) Note: The name S. xfarnhamii was originally based on a hybrid between S. leucophylla (subsp. leucophylla) and S. rubra subsp. rubra. The plant Mellichamp (2008) described as S. x bellii (=S. leucophylla x S. rubra subsp.gulfensis) differs fromS. xfarnhamii in the strict sense atthe rankof subspecies. Parents: (S. leucophylla x S. rubra subsp. gulfensis). Volume 44 March 2015 7 Type locality: West side ofHwy. 87just north ofYellow River, Santa Rosa County, FL, 2008. Holotype: Mellichamp s.n. (UNCC). Similar to those ofits S. leucophylla parent, this hybrid’s leaves exhibit white areoles about the peristome andoperculum. The opercula arewavymarginedandlongerthanwide. The narrowoper- culum is a characteristic ofits S. rubra subsp. gulfensisparent. Somewhat similar, the leaves ofS. x farnhamiinothosubsp. belliiare consistentlylonger(to42cm) thanthoseofS. x readei(Mellichamp 2008).Althoughthe authorsknowofno collections withflowers, the sepals andpetals are likelyred (as in bothparents) andprobably intermediate in size. This is apparently the only site (alongpower lines)wherethisrarehybridhasbeencollected.Atthis site,the seniorauthorin2014 confirmedthat S. xfarnhamii nothosubsp. bellii is still present. S. xformosaVeitch ex Masters Parents: (S. minorWalter x S.psittacina). Type locality: Moistpine barren, nearFitzgerald, Irwin County, GA, 1904. Neotype: Harper2211 (US). Leaves ofthis hybrid are semi-decumbent with bonnet-shaped opercula and translucent areoles. Flowers are distinctive in that the sepals are primarily green and the petals are mostly red. The neotype may be viewed online. Originally named for a horticulturally derivedplant in 1811, it was not until 1904 that a natural hybrid was collected. Since that time, additional collections have been made from southeastern GA. Collections also have been made in Nassau and Baker Counties in northeastern FL. Although previously not reported from Duval County, FL, the senior author ob- servedthishybridinbloomontheUniversityofNorthFloridacampus.Theseplantsarenotrareand are most easily detected inthe field by theirbicolor blooms. S. x gilpiniiBell & Case Parents: (S.psittacina x S. rubra subsp. gulfensis). Type locality: Savanna north ofYellow River and east ofFlorida highway 87; Santa Rosa County, FL, 1955. Holotype: Bell 1523 (NCU). This hybrid produces a rosette ofsemi-decumbent leaves; each bears a bonnet-shaped operculum withtranslucentareoles. Flowers were describedbyBell andCase (1956) as small, faintlysweet-scent- ed, with maroon sepals and petals. Attempts to locate the holotype (or any other naturally-collected A specimens) were unsuccessful. photograph, probably from the original collection, is included in the paper by Bell and Case (1956). According to Mellichamp (2008) this specimen was collected in the samebogwithS. xfarnhamiinothosubsp. bellii(see above).Althoughthe seniorauthordidmanageto findS. xfarnhamiinothosubsp. belliitherein2014 (seeabove), hefoundnoindividualsofS. xgilpinii. Therefore, this hybridwould appearto be extremelyrare ornonexistent inthe wild. Assuming thatthe holotypeislost,thenaneotype shouldbedesignated; however,wewereunabletofindanyotherherbar- ium collections. Designating the above mentionedphotograph, as the neotype wouldbe allowedunder thearticle9.6oftheICBN(McNeiletal.2011).Differencesinfloweringperiodbetweenthetwoparents andtherestrictedrangeofS. rubra subsp.gulfensisarefactorsthatcontributetotherarityofthishybrid. S. x harperiC.R.Bell Parents: S.flava x S. minor). ( Type locality: Sandy bog inpine-barrens nearBloys, Bulloch County, GA, 1901. Neotype: Harper 855 (US). 8 Carnivorous Plant Newsletter Leaves are erect. Exhibiting a darkened splotch near its base, each operculum is revolute. Inter- estingly, the prominent translucent areoles ofits S. minorparent are absent. Flowers unsurprisingly GA exhibit the green sepals and yellow petals ofboth its parents. Only a few collections from and SC were found during this study. Although the parents are sympatric over a large area ofnorthern FL east ofthe Chattahoochee-Apalachicola River system, it is curious that there are no known col- lections from that state. The neotype can be viewed online. S. x mitchellianaW.Bull. Parents: (S. leucophylla x S. rosea). Type locality: Bog 2.5 miles west ofFlorida state line by U.S. 90, Baldwin County,AF, 1948. Neotype: Bell 548 (NCU). The leaves ofthisrobusthybrid are semi-decumbent, eachwith a collar-shape operculumthat is pinched in the middle. When young, white areoles about the operculum andperistome are evident. Flowever, as the leafages, it becomes darkpurple withthe areoles becoming obscured. With its red sepals andpink petals, flowers are similar to its S. rosea parent. Flowers are nearly scentless (pers. obs.). Plants have been collected in Baldwin CountyAF, adjacent Escambia County, FF and from a disjunct population in Liberty County, FL. Bell (1952) statedthat Harper (1918) found this plant inWaltonCountyFL butwe couldnot locate a collectionfromthat county. The curious thing about this hybrid is that it is fairly common in Escambia County, FF and near its type locality inAF, but it seems to be rare or nonexistent in other areas where its parents are sympatric. The senior author observedplants at several locations inEscambiaCounty, FF and some withinthe city limits ofPen- sacolain2014. Considerablevariationinleafshape, colorandsize,probablydueto introgression, is evident inplants that inhabit SplinterHill Bog, Baldwin County,AF (pers. obs.). S. x mooreiMoore ex Masters Parents: (S.flava x S. leucophylla). Type locality: Sandybog southeast ofAmericus, Sumter County, GA, 1901. Neotype: Harper 1021 (US). Marked with red veins, leaves are erect with white areoles about the peristome and operculum. Sepals are green and suffusedwithpale red; the petals are likewise pale red. This commonhybrid’s range extends from southern AF to northwestern FF and southwestern GA. Because its parents freelyhybridize andintrogressionappears common, amyriadofmorphologicalformsareproduced. In2013 and2014, the senior author observedmuch variation in color andvenationpatterns among individuals ofthe Garcon Point peninsula, Santa Rosa County, FF. In 2013, this hybrid was col- lected for the first time in Escambia County, FF by the senior author in the type locality ofS. x naczii. The neotype can be viewed online. Vegetatively similar, S. x areolata also produces a broad spectrum ofvariants. S. x nacziiMellichamp Parents: (S.flava x S. rosea). Type locality: South side ofO.C. Phillips Rd., ca 3 miles east ofPerdido River, Escambia County, FF, 2007. Holotype: Mellichamp s.n. (UNCC). The leaves ofthis robusthybrid are semi-decumbent, eachwith acollar-shape operculumthat is pinched inthe middle. Heavily markedwithred orpurple veins, each leafbears a darkened splotch at the base ofits peristome like that ofits S.flava parent. Flowers are typically pale yellow with Volume 44 March 2015 9 a pink cast (Mellichamp 2008). In 2014, the senior author observed plants in the type locality in Escambia County, FL and in a second location inWalton County, FL. S. xpopeiMasters Parents: S.flava x S. rubra subsp. rubra). ( Type locality: Carthage, Moore County, NC, 1926. Neotype: Harriots.n. (PH). Erect leaves are light green with maroon markings. Earge flowers exhibit maroon sepals with maroonpetals thatare edgedwithyellow(Bell 1952).An illustrationofthe flowerandleafis shown inBell’s (1952) paper. Although they are sympatric over a large range, mostly in the Carolinas, the twoparent speciesproduce few, ifany, natural hybrids. Indeed, there areno otherknownnaturalhy- bridcollectionsbetweenS.flava andanyothermemberoftheS. rubra complex.Neithertheneotype nor any othercollection ofthis taxon was located. S. x readeiBell Parents: (S. leucophylla x S. alabamensis subsp. wherryi). Type locality: DeerPark, Washington County,AL. 1913. Lectotype: Reade 5789 (PENN). Prominentlymarkedwiththe white areoles from its S. leucophylla parent, the leaves ofthisrare hybrid are slender and reddened like those ofits otherparent S. alabamensis subsp. wherryi. Flow- ers are maroon and intermediate in size (Bell & Case 1956). The lectotype was located in the PH herbarium. Only two other collections known to the authors (Bell 1496 and Case P-63) were not located. Due to the restrictedrange ofS. alabamensis subsp. wherryi, this hybrid is rare. S. x rehderiC.R.Bell Parents: (S. minor x S. rubra subsp. rubra). Type locality: Damp savannahs about 12 miles southwest of Shallottee, Brunswick County, NC, 1952. Neotype: Rchdcrs.n. (NCU). Withreddishveins, erectleaves aregreenwithwhite areoles abouttheperistomeandoperculum. Similarto those found in S. x harperi, the opercula are slightly revolute. Flowers are fragrant. Red sepals and petals are edged with yellow (Bell 1952). Bell reported that this hybrid was first col- lectedbyWherry in 1923 inCandlerCounty, GA butno herbarium specimencouldbe located. The neotype was collected in 1951 and consists only ofleaves; however, a black andwhite photograph ofthe plant in flower is attached. An image ofthe neotype is available online. Plants also have been collected inBarnwell and Colleton Counties, SC. S. x swaniana Bull exW.Robinson Parents: (S. minor x S.purpurea subsp. venosa). Type locality: From damp savanna about 12 miles southwest ofShallotte, Brunswick County, NC, 1952. Neotype: Rehders.n. (NCU). Typical ofotherhybrids fromthe S.purpurea complex, leaves are semi-decumbentwith spread- A ing, collar-shaped opercula. few translucent areoles about the base ofthe operculum are present (Bell 1952). Green sepals are suffusedwithred; petals are completelyred. Firstmentionedas ahor- ticulturallyderivedplant in 1887, itwasnotuntil Eliotand Stauffer’s (1951)publicationthatanatu- 10 Carnivorous Plant Newsletter ral hybridwas notedinthe literature. Plants also have beencollectedinBladen County, NC (Bell & Case 1956). The few collections ofthis plant suggest its rarity. The neotype can be viewed online. S. x wrigleyana S.G. Parents: (S. leucophylla x S.psittacina). Type locality: Fowl RiverRoad, south ofTheodore, Mobile County,AL, 1954. Neotype: Case P62 (NCU). The leaves ofthis rare hybrid are semi-decumbent; eachbears a bonnet-shaped operculum with translucent areoles. Like both its parents, the sepals and petals are red; however, the flowers are intermediate in size. Thisplantwas first collectedbyWherrynearFruitdale inWashington, County, AL in 1932 (Bell & Case 1956); however, this collection could not be located. The neotype, which can be viewed online, originally consisted ofleaves only. However, a flower was added later. The flower is from the original specimen that bloomed at the University of Michigan greenhouse in 1955. Bell and Case (1956) also listed a specimen from Escambia County, FL; however, that speci- mencouldnotbe located.Anadditional specimenfromBayCounty, FL was locatedatFLAS 97441 (Beckner et al. 1506). Differences in flowering period between the two parents appear to be the major impediment inproducing this hybrid. Conclusion From this study it is clear that some natural hybrids ofSarracenia are common and others are rare. Judgingbythe largenumbers ofhorticulturally engineeredhybrids, there doesnotappeartobe anygenetic incompatibility among the species ofSarracenia withrespectto theirabilitytoproduce fertile hybrids. Indeed, hybrid swarms are common inthe field (Ellison etal. 2014). Natural hybrids occurinnumbers whenthe two sympatric species are commonandhave at least some overlap intheirrespectiveperiods offlowering. Examples includeS. x areolata, S. xformosa, S. x mitchelliana, andS. x moorei. Conversely, whenthese conditions are notmet, natural hybrids arereduced orabsent. For exam- ple, atemporal difference infloralperiodbetweenparents fostersreproductive isolation. Sarracenia x wrigleyanaprovides agoodexample. Specifically, althoughS. leucophylla andS.psittacina occur sympatrically over a large range, hybrids seldom are produced. In situations where one ofthe par- ents occupies a small range, natural hybrids are likewise, restricted. For example, the rarity ofboth S. x caseiandS. xfarnhamiinothosubsp. belliiis duetothe limitedrange oftheirrespectiveparents S. alabamenis subsp. wherryi and S. rubra subsp. gulfensis. When floral periods between the two parents typically do not coincide and one ofthe parents has a restricted range, then the probability ofprogenybetweenthe two is rare. Such is the case ofS. x gilpinii. Interms oftheirbiological impact, rare Sarracenia hybrids are little more than curiosities. That is, their presence has minimal or no impact on the evolutionary trajectory ofplants in their local group. However, in cases where hybrids are common, the evolutionary impact, at leastwithrespect tothe local group, canbe substantial.Althoughtherehasbeenlittleornomolecularverification, itis likely, based on observable morphological variation, that introgression, resulting inhybrid swarms, occursundertheseconditions.Asmentionedpreviously, hybridswarmsappearto bepresent, atleast in some local groups, where hybrids are common. Acknowledgements: We thank the following for their assistance in locating requested herbarium specimens: Kent Perkins (FLAS); Layne Huiet (CHARL); Shanna Oberriter, Carol McCormick Volume 44 March 2015 11 (NCU); Alina Freire-Fierro (PH). Jan Schlauer provided detailed advice on taxonomy. No special permits were required to visitthe sites described inthis paper. References A Bell, C.R. 1949. cytotaxonomic studyofthe Sarraceniaceae ofNorthAmerica. J. Elisha Mitchell Sci. Soc. 65: 137-166. Bell, C.R. 1952. Natural hybrids in the genus Sarracenia L. History, distribution and taxonomy. J. Elisha Mitchell Sci. Soc. 68: 55-80. Bell, C.R., and Case, F.W.1956. Natural hybrids inthe genus Sarracenia. II Currentnotes on distri- bution. J. Elisha Mitchell Sci. Soc. 72: 142-152. Case, F.W., and Case, R.B. 1976. The Sarracenia rubra complex. Rhodora 78: 270-325. A Elliott, S. 1824. Sketchofthe BotanyofSouth-Carolina andGeorgia. 2:11. Schenck, Charleston, SC. Ellison, A.M., Butler, E.D., Hicks, E.J., Naczi, R.F.C., Calie, P.J., Bell, C.D., and Davis, C.C. 2012. Phylogeny and biogeography ofthe carnivorous plant family Sarraceniaceae. PLoS ONE 7(6): e39291. Ellison,A.M., Davis, C.C., Calie, P.J., andNaczi, R.F.C. 2014. Pitcherplants (Sarracenia) provide a 21st-centuryperspective on infraspecific ranks and interspecific hybrids: amodestproposal* for appropriate recognition andusage. Systematic Botany 39(3): 939-949. Eliot, R., and Stauffer. M. 1951. June in savanna land. WildFlower27: 32-41. Harper, R.M. 1918.TheAmericanPitcherplants.TheElishaMitchell Science Society. 34: 1 10-135. McNeil. J., Barrie, F.R., Buck,W.R., Demoulin,V, Greuter,W., Hawksworth, D.L., Herendeen, PS., Knapp, S., Marhold, K., Prado, J., Prud’homme van Reine, W.F., Smith, G.F., Wiersema, J.H. eds. 2011. International Code ofNomenclature for algae, fungi, and plants (Melbourne Code), Adopted by the Eighteenth International Botanical Congress, Melbourne Australia, July 2011 (electronic ed.). Bratislava, InternationalAssociation forPlantTaxonomy. McPherson, S., and Schnell, D.E. 2011. Sarraceniaceae ofNorthAmerica. Dorset: RedfernNatural History Productions. Mellichamp, T.L. 2008. New names for natural hybrids in Sarracenia. Carniv. PI. Newslett. 37: 112-117. Mellichamp, T.L., andCase, F.W. 2009. Sarraceniaceae. In: Flora ofNorthAmericaEditorial Com- mittee eds. Flora ofNorth America North of Mexico Vol. 8. Magnoliophyta: Paeoniaceae to Ericaceae. OxfordUniversity Press, NewYork and Oxford. 348-363. Neyland R., and Merchant, M. 2006. Systematic relationships ofSarraceniaceae inferred fromnu- clearribosomal DNA sequences. Madrono 53: 223-232. Sheridan, P, and Scholl, B. 1993. Noteworthy Sarracenia collections. Carniv. PI. Newslett. 22: 58- 61. Thiers, B. 2014. IndexHerbariorum:Aglobal directoryofpublicherbariaandassociatedstaff.New YorkBotanical Garden’sVirtual Herbarium, http://sweetgum.nybg.org/ih/. USDA, NRCS. 2014. The PLANTS Database National Plant Data Team, Greensboro, NC 27401- 4901 USA. http://plants.usda.gov/java/.Accessed 11 Sep 2013. 12 Carnivorous Plant Newsletter

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