serpocaulon (polypodiaceae), a new genus segregated from...

INTRODUCTIONThe classical Linnaean genus Polypodium as adopt-

ed in most recent floras and taxonomic treatises com-prises about 125ndash150 species most of them New WorldSometimes additional genera are subsumed withinPolypodium eg Microgramma Phlebodium and Pleo-peltis in the New World and Goniophlebium in the OldWorld (as by Hennipman amp al in Kubitzki 1990) inwhich case Polypodium exceeds 250 species Within abroadly defined morphologically extremely diversePolypodium there are many problematic groups Recentmolecular evidence suggests that Polypodium sl is apolyphyletic assemblage even if several segregate gen-era (eg Microgramma Pecluma and Campyloneurum)are excluded (as in most recent tropical floras eg Try-

on amp Stolze 1993 Moran in Davidse amp al 1995 Mic-kel amp Smith 2004) and that generic boundaries in thiscomplex of genera (Polypodium sl) require substantialrevision (Schneider amp al 2004b)

Within Polypodium sl (as applied in floras and tax-onomic compendia by most authors today) severalgroups have been considered as ldquonaturalrdquo informallyrecognized and taxonomically treated These include 1)Polypodium ss a group of largely north-temperate spe-cies allied to P vulgare L (Haufler amp al 1995 Haufleramp Ranker 1995) with a closely related subgroup P ple-siosorum and allies (Tejero-Diacuteez amp Pacheco 2004Group 1 part of Tryon amp Tryon 1982) in Mexico andCentral America 2) P subpetiolatum and allies (revisedby Maxon 1903 Group 1 part of Tryon amp Tryon1982) 3) the P furfuraceum and P plebeium groups

919

Smith amp al bull Serpocaulon a new fern genus55 (4) bull November 2006 919ndash930

Serpocaulon (Polypodiaceae) a new genus segregated from Polypodium

Alan R Smith1 Hans-Peter Kreier2 Christopher H Haufler3 Tom A Ranker4 amp HaraldSchneider2

1 University Herbarium 1001 Valley Life Sciences Bldg University of California Berkeley California 94720-2465 USA arsmithberkeleyedu (author for correspondence)

2 Albrecht-von-Haller-Institut fuumlr Pflanzenwissenschaften Abt Systematische Botanik Georg-August-Universitaumlt Goumlttingen Untere Karspuumlle 2 37073 Goumlttingen Germany

3 Department of Ecology and Evolutionary Biology University of Kansas Lawrence Kansas 66045-2106 USA4 University Museum and Department of Ecology and Evolutionary Biology University of Colorado Boulder

Colorado 80309 USA

We describe the neotropical fern genus Serpocaulon (Polypodiaceae) segregated from Polypodium and makenew combinations for the forty known species Serpocaulon has been recognized previously as a putativelynatural group by several authors based on morphological evidence but never accorded generic rank Themonophyly of Serpocaulon is strongly supported in chloroplast DNA-based phylogenetic analyses using thegenes rbcL rps4 rps4-trnS intergenic spacer and trnL-F intergenic spacer on 29 samples from 22 speciesSerpocaulon is readily separated from other subclades within Polypodiaceae and especially from Polypodiumwhich is less closely related to Serpocaulon than are several other genera Serpocaulon differs from other NewWorld polypods in the combination of usually long-creeping sparingly branched rhizomes with clathrate(sometimes strongly so) usually peltately attached scales regularly anastomosing (goniophlebioid) veins withindividual areoles chevron-shaped and each with a single free included excurrent veinlet and non-pa-raphysate sori in one to 10 rows between costae and pinna margins In Polypodium the rhizome scales are notclathrate plusmn concolorous and invaginated at the base with somewhat overlapping basal lobes and rhizomesare generally shorter-creeping and more frequently branched the veins are free or forming a single row of are-oles and the sori uniseriate on each side of the costae Most species of true Polypodium are north-temperateMexican or Mesoamerican in distribution while Serpocaulon is entirely neotropical or subtropical with thegreatest number of species in South America There appears to be no convenient infrageneric division of thespecies within Serpocaulon although the most widespread species S triseriale is somewhat isolated at thebase of the clade Species with the thinnest rhizomes also have the fewest scales and generally grow at highelevations Species having the scaliest rhizomes with scales spreading and strongly overlapping form a weak-ly supported subclade as do species with multiseriate rows of areoles between costae and pinna margins

KEYWORDS ferns phylogeny Polypodiaceae rbcL rps4 rps4-trnS IGS Serpocaulon trnL-F IGS

(Maxon 1916a 21 spp Groups 4 and 6 parts of Tryonamp Tryon 1982) 4) P polypodioides (Maxon 1916bWeatherby 1939 Group 4 part of Tryon amp Tryon1982) 5) P squamatum and relatives (Maxon 1916bWeatherby 1947 revised by de la Sota 1966 22 sppGroup 5 part of Tryon amp Tryon 1982) 6) the P thys-sanolepis group (Wendt 1980 Group 4 part of Tryon ampTryon 1982) and 7) P loriceum P fraxinifolium andallies an entirely neotropical group of about 40 specieswith regularly anastomosing (goniophlebioid) veins andoften long-creeping cordlike rhizomes bearing partiallyor entirely clathrate scales (most of Group 2 of Tryon ampTryon 1982 species taxonomy 23 spp revised byHensen 1990) Because nearly all of the major sub-groups of Polypodium sl occur in Mexico and CentralAmerica and these areas are a center of diversity for sev-eral of the subgroups many species have been treated infloristic contexts by Lellinger (1989) Moran (in Davidseamp al 1995) and by Mickel amp Smith (2004) Howeverthe species allied to P loriceum and P fraxinifolium arepredominantly South American and are less well studiedexcept for work by Hensen (1990) which was followedin the main in the floristic account for Peru by Tryon ampStolze (1993)

Existing evidence (Schneider amp al 2004b) supportsthe placement of species in 3) through 6) as delineatedabove in Pleopeltis All of these groups contain speciesthat bear scales on the abaxial lamina A few scaly-blad-ed species of Polypodium have already been transferredto Pleopeltis (Windham 1993) as precursor to theirtreatment in the Flora of North America North of Mexico(Andrews amp Windham in FNA Ed Comm 1993) how-ever most scaly species remain in Polypodium to betransferred to Pleopeltis elsewhere (Smith amp al inprep) Species now placed in the segregate generaDicranoglossum Neurodium Microphlebodium andPseudocolysis also appear to be closely allied to and per-haps inseparable from Pleopeltis sl In the present paperwe address the species mentioned under 7) above andthese lack a name at generic rank This new genus wehere name Serpocaulon alluding to the long-creepingsnakelike rhizomes found in nearly all species

Many of the species of Serpocaulon were initiallydescribed in Goniophlebium (Blume) C Presl (seeAppendix 2 electronic supplement) a genus of about 25species occurring from India throughout southeast andeastern Asia the Himalayan region and Malesia toAustralasia Melanesia Samoa and Micronesia (Roumldl-Linder 1990) The name Goniophlebium is derived fromthe Greek gonia meaning angle and phleps vein refer-ring to the characteristic venation pattern formed by reg-ularly anastomosing veins that often form a succession ofchevron-shaped areoles between the pinna midribs andmargins Although the paleotropical species of Gonio-

phlebium and the neotropical species of Serpocaulonhave similar venation and both genera bear clathrate rhi-zome scales the similarities are convergent as indicatedby their positions in phylogenetic trees based on molec-ular data (see Schneider amp al 2004b) Goniophlebiumss falls within an almost exclusively monophyletic OldWorld clade (as far as known the lone exception is a sin-gle disjunct species of Platycerium in the AndesSchneider amp al 2004b) whereas Serpocaulon nests in apredominantly New World clade (with a few African out-liers) containing the genera Microgramma Campyloneu-rum Pleopeltis Pecluma and Polypodium ss as wellas the pantropical grammitid clade Roumldl-Linder (1990)and Lellinger (1993) also specifically excluded all NewWorld species of Polypodium from Goniophlebium onthe basis of several morphological characters eg theabsence of black sclerenchyma strands in the rhizomespeltate-based rhizome scales the absence of hairy orscaly paraphyses in the sori and the folded perine aroundthe spores

In this paper we restrict the application ofPolypodium L (Sp pl 2 1082 1753) with lectotype Pvulgare L (chosen by John Smith Hist fil 88 1875) toa group of predominantly temperate (mainly north-tem-perate) species plus a less widespread tropical and sub-tropical group of Mexican-Mesoamerican speciesPolypodium as defined in this narrow sense then com-prises about 15 species in temperate regions of AmericaEurope Asia and Africa (Haufler amp al 1995 Haufler ampRanker 1995) and perhaps another 15 species in Mexicoand Mesoamerica The species have deeply pinnatifid orweakly 1-pinnate blades concolorous or sometimesslightly bicolorous orange-tan to brown non-clathraterhizome scales free or weakly anastomosing venation(each areole with a single free included veinlet) soriterminal on ends of free included veinlets channeledstipes filamentous or branched paraphyses and morefrequently branched rhizomes

Within Serpocaulon eight (of about 40 total) specieshave been included previously (as Polypodium) in a phy-logenetic study using the chloroplast DNA regions rbcLrps4 and rps4-trnS intergenic spacer (Schneider amp al2004b) Polypodium adnatum Kunze ex Klotzsch[voucher re-identified herein as S richardii (Klotzsch)Domin] P fraxinifolium Jacq P funckii Mett (voucherre-identified as S ptilorhizon (H Christ) A R Sm) butnot used in the current study) P levigatum Cav Ppatentissimum Mett ex Kuhn P ptilorhizon H Christ Ptriseriale Sw and P wiesbaueri Sodiro [= Peleutherophlebium (Feacutee) Mett in this paper] In a maxi-mum likelihood analysis (Schneider amp al 2004b) thisclade is monophyletic with clade confidence values of100 (Bayesian posterior probability PP) and non-para-metric bootstrap support of 99 In the Schneider amp al

Smith amp al bull Serpocaulon a new fern genus 55 (4) bull November 2006 919ndash930

920

analysis this clade is weakly supported (Bayesian PP052) as sister to the entire pantropical grammitid clade(Grammitidaceae in many classifications) comprisingabout 750 species In molecular analyses Serpocaulon iswell removed from other species groups currently treat-ed in Polypodium namely the scaly species (includingPleopeltis Humb amp Bonpl ex Willd ss MarginariaBory Polypodium furfuraceum and allies P myriolepisand allies P guttatum and allies Dicranoglossum J SmMicrophlebodium L D Goacutemez (Goacutemez 1985) Pseudo-colysis L D Goacutemez (Goacutemez 1977) and possibly sever-al other segregate genera) as well as from thePolypodium vulgare group and from Pecluma M GPrice In the present study we developed and analyzednew molecular data from 15 additional species as well asincorporating examples from previously sequencedspecies for a total of 22 species now examinedmdashrepre-senting slightly more than half of the currently knownspecies

It is the objective of this paper to present the molec-ular and morphological evidence for characterizingSerpocaulon formally name the new genus and effecttransfer of species belonging to it

MATERIALS AND METHODSSequences of four chloroplast DNA regions rbcL

rps4 rps4-trnS intergenic spacer and trnL-F intergenicspacer were generated using primers and protocols asdescribed in previous studies (Haufler amp Ranker 1995Schneider amp al 2002 2004a b Haufler amp al 2003Janssen amp Schneider 2005) With the exception of newlygenerated sequences for samples of Serpocaulon andtrnL-F for Microgramma percussa all sequences used inour current study were published in previous papers(Schneider amp al 2004a b) The taxon dataset in the cur-rent paper is a subset of the one used by Schneider amp al(2004a) plus additional species of SerpocaulonAppendix 1 provides the GenBank accession numbersfor all samples of Serpocaulon including previouslypublished sequences For information concerningsequences of taxa not belonging to Serpocaulon exceptfor Microgramma percussa see Schneider amp al (2004a)All sequences were aligned manually using MacClade40 (Maddison amp Maddison 2000) Ambiguously align-ing regions were excluded from all analyses

Phylogenetic analyses were performed using PAUP40b10 (Swofford 2000) Dictymia brownii (Wikstr)Copel was chosen as the outgroup representing the lox-ogrammoid lineage which has been shown to be the sis-ter lineage of all other polygrammoid ferns (Schneider ampal 2004b) Maximum parsimony analyses were generat-ed using the heuristic search mode with TBR branch

swapping 1000 random addition sequence replicatesand MULTREES on Bootstrap values were calculatedwith 10000 bootstrap replicates using the heuristicsearch mode with TBR branch swapping and 100 randomaddition sequence replicates The absence of incompati-bilities among the four chloroplast DNA regions wascontrolled by visual comparison of the results of boot-strap analyses of each of the four cpDNA regionsDifferences were judged to be inconsequential and thefour regions were combined in all analyses presentedMaximum likelihood analyses were carried out withmodels selected using the hierarchical likelihood test asimplemented in Modeltest (Posada amp Crandall 1998)The same model GTR + I + Γ was selected using theAIC criterion in Modeltest It was implemented with thefollowing parameters A = 02974 C = 01980 G =02020 T = 02944 A-C = 12135 A-G = 34737 A-T =04586 C=G = 10138 C-T = 43370 G-T = 1000 I =02371 and Γ = 0999 in the files executed in PAUP40b10 Tree searches were performed using the heuristicmode with TBR branch swapping and 100 random addi-tion sequence replications

Bayesian inference of phylogeny was carried outwith MrBayes 30 (Huelsenbeck amp Ronquist 2001) withthe GTR + I + Γ model implemented for 4 chains with10000000 generations of which each 1000th generationwas sampled The burn-in phase was identified visuallyand excluded before inferring Bayesian posterior proba-bilities

RESULTSMaximum likelihood analysis of the combined

dataset yielded an optimal tree with a -lnL = 02023397(Fig 1) Maximum parsimony analysis of the same data-set resulted in 348 most parsimonious trees with a lengthof 3023 steps consistency index of 04408 homoplasyindex of 05592 retention index of 07033 and arescaled consistency index of 03899 (Fig 2)

Serpocaulon was monophyletic in maximum likeli-hood (ML) maximum parsimony (MP) and BayesianInference of phylogeny (BY) with strong support in BYanalysis (Fig 1 PP = 100) and MP analysis (Fig 2 BS= 99) Serpocaulon was not closely related to othersegregates of Polypodium such as GoniophlebiumPleopeltis Pecluma and Polypodium ss Serpocaulontriseriale was strongly supported as sister to the remain-ing 21 species of Serpocaulon included in this study(Figs 1 2) In the ML analysis S sessilifolium (clade IIin Figs 1 and 2) was sister to the remaining 20 speciesThese 20 species clustered into two clades (clade III + IVin Fig 1) One clade included the species S lasiopus toS silvulae (clade III in Fig 1) whereas the other includ-


921

ed S eleutherophlebium to S maritimum (clade IV inFig 1) The latter clade had strong support in the BYanalysis (PP = 100) and poor bootstrap support in theMP analysis (BS = 72) The clade including S lasiopusto S silvulae was not monophyletic in the strict consen-sus tree of the 348 most parsimonious trees (clade III inFig 2) but it was present in 95 of these trees In the

MP analysis S sessilifolium (clade II in Fig 1) was sis-ter to the clade comprising S eleutherophlebium to Smaritimum (clade IV in Fig 1)

All five species represented by more than one speci-men formed clades at least in the ML analysis The fourspecimens of S sessilifolium formed a strongly support-ed clade (BS = 100 Bayesian PP = 100) Two speci-mens of S subandinum were sister to each other withbootstrap support of 97 and PP = 100 Two specimensof S richardii also formed a clade in the ML tree but thisclade was found in only 74 of the MP trees A BayesianPP of 098 supported the monophyly of this species Twospecimens of S crystalloneuron formed a clade with Ssilvulae in the ML tree with PP of 100 but only 95 ofthe most parsimonious trees possessed this clade Thetwo specimens of S intricatum formed a clade with Slatipes and S latissimum in both ML and MP trees Thisresult was supported by a Bayesian PP of 100 and abootstrap value of 90


001 substitutionssite

Dictymia brownii

Aglaomorpha acuminataAglaomorpha coronans

Aglaomorpha meyenianaDrynaria rigidula

Arthromeris lehmannii

Selliguea enervis

Selliguea heterocarpa

Selliguea feeiSelliguea lanceolata

Selliguea triloba

Thylacopteris papillosa

Goniophlebium Lemmaphyllum carnosum

Leptochilus macrophyllusMicrosorum cuspidatum

Microsorum musifoliumMicrosorum punctatum

Microsorum linguiforme

Lecanopteris carnosaLecanopteris sinuosua

Pyrrosia Platycerium

Microgramma percussaCampyloneurum angustifoliumCampyloneurum chlorolepis

Terpsichore Adenophorus oahuensis

Pecluma ptilodon

Pleopeltis angustaPleopeltis polypodioides

Polypodium pellucidumPolypodium vulgare sl

Polypodium roseiPolypodium guttatum

S triseriale Jimenez 1994

S attenuatum Moreno 10406S gilliesii Ehrich 314

S vacillans Mexia 5856a

S lasiopus Lewis 35186S wagneri Smith 1171

S crystalloneurum Jimenez 1176S crystalloneurum Jimenez 1109

S silvulae Jimenez 1093

S sessilifolium Jimenez 1034

S sessilifolium Jimenez 755S sessilifolium Jimenez 976

S levigatum Jimenez 756

S maritimum Smith sn

S patentissimum Smith 2829

S eleutherophlebium Smith 2877

S richardii Moran 6258S richardii Jimenez 1986

S appressum Jimenez 862S fraxinifolium Jimenez 696

S giganteum Jimenez 1035

S intricatum Jimenez 1324S latissimum Jimenez 813

S latipes Bach 1305S intricatum Jimenez 694

S subandinum Hutchinson 5872S subandinum Moran 6858

S ptilorhizon Jimenez 1102


I

II

III

IV

Fig 1 Phylogram with the highest likelihood value foundin the maximum likelihood analysis of the combineddataset Thick lines indicate branches with Bayesian pos-terior probabilities of ge095 The arrow indicates theSerpocaulon clade Vertical lines on the right indicatefour clades within Serpocaulon discussed in the resultsAsterisks indicate composite taxa that are based onsequences generated from two species of an unambigu-ously monophyletic genus or lineage (see Schneider ampal 2004a for further information) All samples of Serpo-caulon are given with species name and the correspon-ding voucher

Dictymia brownii

Aglaomorpha acuminataAglaomorpha coronansAglaomorpha meyenianaDrynaria rigidula


Selliguea enervis



Selliguea triloba



Leptochilus macrophyllusMicrosorum cuspidatumMicrosorum musifoliumMicrosorum punctatum






Pecluma ptilodon








S crystalloneurum Jimenez 1176 S crystalloneurum Jimenez 1109S silvulae Jimenez 1093S sessilifolium Jimenez 1034










S latipes Bach 1305S intricatum Jimenez 694S subandinum Hutchinson 5872S subandinum Moran 6858



100

100

100

8493

98

76

77

100

100

100

100

100

100

9290

98100

97

99

100100

93

91

72

90

I

II

III

IV

75

Fig 2 Strict consensus tree of 348 most parsimonioustrees obtained in a maximum parsimony analysis of thecombined dataset Bootstrap values gt70 are givenabove branches For further information see Fig 1

922

DISCUSSIONThe monophyly of Serpocaulon is strongly support-

ed in all analyses of the cpDNA sequence data data thatinclude slightly more than half of the currently acceptedspecies In all phylogenetic analyses the lineage isclearly separated from other lineages traditionallyassigned to Polypodium sl (Schneider amp al 2004a b)and it is unambiguous whether a given species ofPolypodium sl is included or not Serpocaulon appearsmorphologically well-defined and has been recognizedpreviously as a putative natural group by various authors(Tryon amp Tryon 1982 Group 2 Hensen 1990)although Tryon and Tryon included three species thatboth we and Hensen exclude Polypodium decumanumWilld [= Phlebodium decumanum (Willd) J Sm] Pdecurrens Raddi [= Campyloneurum decurrens (Raddi)C Presl] and Polypodium kunzeanum C Chr]Lellingerrsquos (1993) definition of Polypodium subg Poly-goniophlebium roughly coincides with our circumscrip-tion of Serpocaulon except that we exclude Polypodiumplectolepidioides Rosenst and P rhachypterygiumLiebm (which may be more closely related toPecluma) as well as P plesiosorum Kunze (which weretain in a redefined and smaller Polypodium seeTejero-Diacuteez amp Pacheco 2004) all three of these specieswere mentioned by Lellinger as belonging to subgPolygoniophlebium

Hensen (1990) attempted a revision of the Poly-podium loriceum group (our genus Serpocaulon) basedon morphological characters and several new specieshave been described since (Kessler amp Smith 2005)Because Hensenrsquos (1990) circumscription of the ldquolorice-umrdquo group is basically the same as ours it is of interestto compare the results of his morphology-based cladisticanalysis with our molecular results With Hensenrsquosdataset homoplasy was acknowledged to be a majorproblem A further problem was his selection ofCampyloneurum as outgroup because this genus maynot be the closest relative of Serpocaulon (Schneider ampal 2004b) Hensen assessed 18 characters of the rhi-zomes rhizome scales fronds indument and spores andhis consensus cladogram shows two major lineages oneof which corresponds in part with a clade found in allanalyses of our cpDNA dataset that was strongly sup-ported in BY (PP =100) but poorly supported in MP (BS= 72) (Fig 2 clade IV) This clade includes taxa withless scaly rhizomes with many of the species havingwholly or partly appressed scales Hensenrsquos ldquoscalyrdquo cladealso included S sessilifolium which is either sister to theother major clade (Clade IV) of Serpocaulon (MP) or sis-ter to clades III + IV (ML) based on our cpDNA evi-dence Taking into account the basal position of S trise-riale the cpDNA data suggest a trend from densely scaly

rhizomes towards less dense indument and decreasingsize of the scales in this genus In Hensenrsquos analysis Striseriale was sister to a clade that included taxa withreduced indument density This result may reflect thepresence of many plesiomorphic features in this puta-tively earliest diverging species of Serpocaulon

We see other resemblances of putatively relatedspecies in the Hensen cladogram and our molecular treeThese include 1) a Hensen-clade containing species with1-pinnate blades and 3+ rows of areoles and sori betweencostae and pinna margins (containing S appressum Sfraxinifolium S richardii and two other species notsampled by us S adnatum and S caceresiimdasha subset ofour clade IV in the ML analysis (Fig 1) and 2) a cladecontaining most of the species with densely scaly rhi-zomes (comprising S crystalloneuron S gilliesii Slasiopus S vacillans and S wagneri)mdashbasically ourclade III in the ML analysis (Fig 1) Thus the genealog-ical composition of both Hensenrsquos morphology-basedtree and our molecular tree show similarities althoughsome of the results are difficult to compare because sam-pling in the two studies differed and we have also adopt-ed different circumscription of certain species

Currently evidence suggests some trends in the mor-phological evolution of Serpocaulon Densely scaly rhi-zomes appear to be plesiomorphic with a trend towardreduction in the number and size of scales on the rhi-zomes Narrowing of the rhizome diameter also seems tobe correlated with scale reduction Species with the thin-nest rhizomes (smallest diameter) and fewest scales forma grade at the base of our clade IV in the ML analysis(Fig 1) S eleutherophlebium S ptilorhizon S suban-dinum and S levigatum however S patentissimumwith the fewest rhizome scales and thinnest rhizomes isnot clearly part of this clade Not only is scale numberreduced in these species but the rhizome scales are oftentightly appressed or spreading only at the scale tips theultimate reduction is seen in S patentissimum S leviga-tum and S ptilorhizon where the rhizomes are sub-glabrous these species tend to grow at high elevationsSpecies having dense persistent spreading and stronglyoverlapping scales fall toward the base of the tree in bothML and MP analyses (Figs 1ndash2) S attenuatum S crys-talloneuron S gilliesii S lasiopus S sessilifolium Striseriale and S wagneri Several of these grow at lowelevations and all except S triseriale have a single rowof sori on each side of the costa

Another trend within Serpocaulon appears to beincreasing pinnation (ultimately to completely free non-adnate pinnae) in the clade comprising S maritimum Sfraxinifolium S appressum S giganteum and S richar-dii Several other unsampled species are expected to fallwithin this clade eg S caceresii and S adnatum Thosespecies showing increased pinnation (free non-adnate


923

pinnae) also show a greater number of rows of areoles(and sori) between the costae and pinna margins

Serpocaulon is related to several genera with an ex-clusively or nearly exclusively neotropical distributionWe refrain from drawing conclusions about the closestrelative until denser sampling of the putative closest lin-eages such as Microgramma and the grammitids can beperformed In Schneider amp al (2004b) Serpocaulon wasfound to be sister to grammitids but these relationshipswere poorly supported Further evidence is needed toconfirm this hypothesis or to confirm alternative hypoth-eses such as close relationships between Microgrammaand Serpocaulon or between Campyloneurum and Serpo-caulon Further sampling must also precede explorationof the radiation of Serpocaulon in the Neotropics andpossible correlations between the origin of Andean habi-tats and the diversification of the genus in that areahome of nearly three-fourths of the species in the genusWe note that the basalmost species in the genus S trise-riale has the largest distribution of any species in Serpo-caulon covering nearly the entire distribution of the ge-nus latitudinally (but not elevationally) Biogeographicaland ecological trends can be only generally stated usingexisting data (1) Serpocaulon shows the greatest diver-sity in South America (2) species occurring at lowestelevations in the genus usually below 500 m appear tobe members of the basalmost branches of the tree (Sattenuatum primarily Amazonian S triseriale mostly100ndash1500 m and S wagneri mostly 0ndash1000 m) and (3)those species having thinnest rhizomes and fewest scalesare entirely or predominantly middle- to high-Andean incomparison to many of the other species of Serpocaulonwhich are more widespread in montane forests or main-ly at low to middle elevations

Although the current sampling is insufficient to drawconclusions about the relationships among the species ofSerpocaulon as well as the monophyly of some speciesthis summary illustrates that the sampling is adequate tooutline some issues that need to be considered in futurestudies and in any exhaustive revision of the genus Adenser taxon sampling with additional molecular mark-ers must be obtained before these issues are explored indetail

TAXONOMIC TREATMENTSerpocaulon A R Sm gen nov ndash Type Serpo-

caulon loriceum (L) A R Sm [equiv Polypodium loriceumL] Figs 3 4

Polypodium L subg Polygoniophlebium LellingerAmer Fern J 83 37 1993 ndash Type Polypodium fraxini-folium Jacq [equiv Serpocaulon fraxinifolium (Jacq) ARSm]

A Polypodio sensu stricto venis ordinate anasto-mosantibus areolis cheveroniformibus (similaribusGoniophlebio et Thelypteridi subg Meniscio) 1ndash10-jugatis inter costas et margines pinnarum omnis areolisvenula singulari inclusa excurrenti libero paleis rhizoma-tum clathratis minimum centraliter vel basaliter gener-aliter peltatis rhizomatibus plerumque longirepentibusparce ramosis distributione neotropico differt a Pleopeltisensu lato laminis pinnatisectis vel 1-pinnatis paleis car-entibus abaxialiter et adaxialiter in venis et interveniis dif-fert a Pecluma venis ordinate anastomosantibus differt

Epiphytic or epipetric infrequently terrestrialRhizomes usually very long-creeping short-creeping in afew species sparingly branched the phyllopodia(05)5ndash20 times the rhizome width apart sometimes glau-cous root proliferations absent black sclerenchymastrands absent vascular bundles sometimes with scle-renchyma sheaths scales round oblong lanceolate orlanceolate-attenuate and with a filiform tip bicolorouspart of each scale clathrate with the side walls dark brownor black sometimes the lumina also blackish so that thescales appear completely black generally peltatelyattached (or with strongly overlapping lobes at a deeplycordate base) surfaces glabrous (lacking rhizoidal surfaceindument) margins entire or toothed Stipes stramineousto light brownish nearly terete proximally flattened tochanneled distally Blades pinnatifid to pinnate rarelysimple (S levigatum) or shallowly pinnatifid monomor-phic stipitate articulate on short phyllopodia laminaeglabrous hairy with scales absent or scattered clathratescales present along the costae and a few on the rachisesveins regularly anastomosing (goniophlebioid) areoleschevron-shaped and each with a single free includedvein Sori round or nearly so in one to 10 rows betweencostae and margins exindusiate each sorus terminal on afree included vein paraphyses absent or present andshort 2ndash3-celled glandular not easily seen sporangiaglabrous Spores light yellowish to whitish bilateral exineverrucate perispore relatively thin or occasionally thick-ened generally tuberculate occasionally winged x = 37

Distribution ca 40 species several recentlydescribed (Kessler amp Smith 2005) and several moreneeding description Preponderantly S American butwith 14 species extending or restricted to the AntillesCentral America and southern Mexico (S antillense Sattenuatum S dissimile S eleutherophlebium S fal-caria S fraxinifolium S giganteum S levigatum Sloriceum S loriciforme S maritimum S ptilorhizon Striseriale S wagneri) The only species restricted to theAntilles or to North America north of Colombia are Santillense and S falcaria while 26 species are confinedto South America

Serpocaulon differs from Polypodium in having reg-ularly anastomosing (goniophlebioid) veins with individ-


924


925

Fig 3 Serpocaulon species A S caceresii portion of pinna with multiseriate sori B S dissimile portion of clathratescale C S fraxinifolium glaucous rhizomes with sparse round scales D S dasypleuron portion of pinnatisect bladewith uniseriate sori E S levigatum portions of simple blades F S triseriale fronds with 1-pinnate blades G S frax-inifolium portion of pinna H S dissimile portion of pinnae Photo credits R C Moran httpwwwplantsystematicsorg


Fig 4 Serpocaulon species AndashC S dissimile (Hallberg 1281 NY) A pinnae and rhizome B pinna detail C rhizomescale DndashF S falcaria (Mickel 1527 NY) D pinnae E pinna detail F rhizome scale GndashK S fraxinifolium (Tuumlrckheim642 NY) G pinnae H rhizome J rhizome scale K pinna detail LndashO S triseriale (Hallberg 1284 NY) L pinnae Mpinna detail N rhizome O rhizome scale 3 cm bar scale applies to A D G and L smaller unlabeled bar scales are1 mm and apply to rhizome scales Reproduced with permission from Mickel amp Smith (2004)

926

ual areoles chevron-shaped and each with a single freeincluded veinlet sori are in one to 10 rows betweencostae and margins (Figs 3ndash4) Polypodium has veinsfree casually anastomosing or (especially in Mexicanand Mesoamerican species) with areoles forming a singlerow between costae and pinna margins and species haveonly a single row of sori on each side of the costaeAdditionally the rhizome (and costal scales if present)in Serpocaulon are clathrate (Fig 3B) sometimesstrongly so at least in the central or basal part of thescales although in some species the lumina are alsorather blackish or obscure rhizome scales are also usual-ly peltately attached at their base or with a deeply cordi-form base with strongly overlapping lobes (eg in S ses-silifolium) In Polypodium the rhizome scales are plusmn con-colorous and invaginated at the base with somewhatoverlapping basal lobes Rhizomes of Serpocaulon aregenerally much longer-creeping and less frequentlybranched than in Polypodium (Figs 3C 4A H N)Polypodium ss is largely north-temperate Mexican andMesoamerican while Serpocaulon is tropical or subtrop-ical in distribution with most species in South America

From Pleopeltis Serpocaulon differs in the absenceof scales on veins and tissue between veins and generallyeasily visible venation Many species of Serpocaulonhave glandular or acicular hairs (Figs 3D 4B) which arelacking in most species of Pleopeltis Pecluma differsfrom Serpocaulon in having free or sporadically anasto-mosing veins a single row of sori on each side of thecostae often reduced proximal pinnae non-clathrateoften comose rhizome scales generally short-creepingrhizomes and pinnatisect blades with usually more than30 pairs of pinnae per blade and pinnae usually dilated atthe base Phlebodium differs from Serpocaulon in theshorter-creeping generally much scalier rhizomes non-clathrate reddish to orangish concolorous spreadingscales the vein areoles more irregularly arranged and bythe sori served by two excurrent veinlets that are joined attheir tips Phlebodium is only remotely related to Serpo-caulon in phylogenetic analyses (Schneider amp al 2004b)

Serpocaulon comprises several somewhat ill-definedand informal morphological groups (1) the S loriceumgroup (S dasypleuron S falcaria S intricatum S lati-pes S latissimum S loriceum Figs 3D 4DndashF) withrhizomes glaucous or not rhizome scales scattered (orrhizome nearly naked) to dense clathrate at least cen-trally round to ovate to ovate-lanceolate blades pinnati-sect or with strongly adnate and often surcurrent pinnaveins anastomosing in 1 (occasionally 2 or 3) row(s)scattered scales present on midribs abaxially these darkand clathrate (2) the S fraxinifolium group (S adnatumS appressum S caceresii S fraxinifolium S gigan-teum S richardii Figs 3A C G 4GndashK) with rhizomesnot glaucous scales scattered (or rhizomes nearly naked)

to dense decidedly clathrate (at least centrally) andround to ovate to ovate-attenuate blades 1-pinnate withpinnae sessile or adnate not surcurrent veins regularlyanastomosing in (1ndash)2ndash10 series between midribs(costae) and pinna margins (3) the S subandinum group(S eleutherophlebium S funckii S levigatum S ptilo-rhizon Fig 3E) with narrow sparsely scaly sometimesglaucous rhizomes pinnatisect blades (simple in S levi-gatum) and (4) the S lasiopus group (S catharinae Scrystalloneuron S dissimile S lasiopus S panorenseand S wagneri Figs 3B 4AndashC) with rhizomes denselyscaly scales usually spreading blades pinnatisect or 1-pinnate with adnate pinnae

Chromosome numbers reported for species of Serpo-caulon are diploid 2n = 37 II (or 2n = 74) for most spe-cies a common number in Polypodiaceae Polypodiumptilorhizon and P fraxinifolium (Sorsa 1966)Polypodium chnoodes (Walker 1966) and Polypodiumloriceum and P triseriale (Smith amp Mickel 1977Walker 1985) The tetraploid condition 2n = 74 II hasalso been reported for P triseriale (Walker 1966) x = 37is by far the commonest base number in neotropicalpolypodioid ferns and also occurs in Polypodium ss

The spores of Serpocaulon are typical of other poly-pods light yellowish to whitish ellipsoid or reniformand monolete and with a generally tuberculate perispore(eg Serpocaulon levigatum S latipes S maritimum Smeniscifolium S subandinum and S triseriale Hensen1990) surrounding a low-verrucate exospore (egSerpocaulon latipes Tryon amp Lugardon 1991 344 Swagneri Hensen 1990 fig 62) The illustration of thespores of S sessilifolium by both Hensen (1990) and byTryon and Lugardon (1991 fig 1316) shows a perisporewith loose folds (wings) that appears unlike most othermembers of the genus Serpocaulon wagneri also haslow perispore folds overlying a low-tuberculate pattern

New combinationsSerpocaulon adnatum (Kunze ex Klotzsch) AR Sm

comb nov equiv Polypodium adnatum Kunze exKlotzsch Linnaea 20 395 1847 ndash DistributionColombia Venezuela and Guianas to Bolivia Brazil

Serpocaulon antillense (Maxon) AR Sm comb novequiv Polypodium antillense Maxon Proc Biol SocWash 43 83 1930 ndash Distribution Cuba JamaicaHispaniola Guadeloupe Martinique GuayananVenezuela

Serpocaulon appressum (Copel) AR Sm comb novequiv Polypodium appressum Copel Univ Calif PublBot 10 305 pl 66 1941 ndash Distribution PeruBolivia

Serpocaulon attenuatum (Humb amp Bonpl ex Willd)AR Sm comb nov equiv Polypodium attenuatumHumb amp Bonpl ex Willd Sp pl ed 4 5 191


927

1810 ndash Distribution Nicaragua to Panama Vene-zuela Guianas Colombia to Bolivia Brazil

Serpocaulon caceresii (Sodiro) AR Sm comb nov(Fig 3A) equiv Polypodium caceresii Sodiro Cryptvasc quit 360 1893 ndash Distribution Colombia toBolivia and French Guiana Brazil

Serpocaulon catharinae (Langsd amp Fisch) AR Smcomb nov equiv Polypodium catharinae Langsd ampFisch Pl voy Russes monde 1 t 9 1810 ndashDistribution Brazil

Serpocaulon chacapoyense (Hook) AR Sm combnov equiv Polypodium chacapoyense Hook HookerrsquosIcon 1 t 69 1937 ndash Distribution Peru

Serpocaulon concolorum (M Kessler amp AR Sm) ARSm comb nov equiv Polypodium concolorum MKessler amp AR Sm Candollea 60 277 2005 ndashDistribution Bolivia

Serpocaulon crystalloneuron (Rosenst) AR Smcomb nov equiv Polypodium crystalloneuron RosenstRepert Spec Nov Regni Veg 11 57 1912 ndashDistribution Peru Bolivia

Serpocaulon dasypleuron (Kunze) AR Sm comb nov(Fig 3D) equiv Polypodium dasypleuron Kunze Lin-naea 9 43 1834 ndash Distribution Colombia toBolivia

Serpocaulon dissimile (L) AR Sm comb nov (Figs3B 3H 4AndashC) equiv Polypodium dissimile L SystNat ed 10 2 1325 1759 ndash Distribution SMexico to Ecuador Antilles Trinidad

Serpocaulon eleutherophlebium (Feacutee) AR Sm combnov equiv Goniophlebium eleutherophlebium FeacuteeMeacutem foug 5 255 1852 ndash Distribution Costa RicaPanama Colombia Venezuela Ecuador

Serpocaulon falcaria (Kunze) AR Sm comb nov(Fig 4DndashF) equiv Polypodium falcaria Kunze Linnaea18 316 1844 [1845] ndash Distribution S Mexico toHonduras

Serpocaulon fraxinifolium (Jacq) AR Sm combnov (Figs 3C 3G 4GndashK) equiv Polypodium fraxinifoli-um Jacq Collectanea 3 187 1789 ndash DistributionMexico to Bolivia Guianas S Brazil

Serpocaulon funckii (Mett) AR Sm comb nov equivPolypodium funckii Mett Abh SenckenbergNaturf Ges 2 57 1857 ndash Distribution Venezuelaand Colombia to Bolivia

Serpocaulon giganteum (Desv) AR Sm comb nov equivPolypodium giganteum Desv Meacutem Soc LinnParis 6 236 1827 ndash Distribution Costa Rica toBolivia S Brazil

Serpocaulon gilliesii (C Chr) AR Sm comb nov equivPolypodium gilliesii C Chr Index fil 529 1906 (anom nov for Polypodium pubescens Hook amp Grev(non L 1759) ndash Distribution Peru Bolivia NWArgentina

Serpocaulon intricatum (M Kessler amp AR Sm) ARSm comb nov equiv Polypodium intricatum MKessler amp AR Sm Candollea 60 278 2005 ndashDistribution Ecuador Peru Bolivia

Serpocaulon lasiopus (Klotzsch) AR Sm comb novequiv Polypodium lasiopus Klotzsch Linnaea 20 3931847 ndash Distribution Venezuela Ecuador PeruBolivia

Serpocaulon latipes (Langsd amp Fisch) AR Sm combnov equiv Polypodium latipes Langsd amp Fisch Pl voyRusses monde (Icon Fil) 1 10 t 10 1810 ndashDistribution Bolivia S Brazil

Serpocaulon latissimum (RC Moran amp B Oslashllg) ARSm comb nov equiv Polypodium latissimum RCMoran amp B Oslashllg Nordic J Bot 15 184 1995 ndashDistribution Ecuador Bolivia

Serpocaulon levigatum (Cav) AR Sm comb nov(Fig 3E) equiv Polypodium levigatum Cav Descr Pl244 1801 ndash Distribution Costa Rica GuadeloupeColombia to Guianas and Bolivia

Serpocaulon loriceum (L) AR Sm comb nov equivPolypodium loriceum L Sp Pl 2 1086 1753 ndashDistribution Nicaragua to Panama AntillesTrinidad Colombia Venezuela Guianas

Serpocaulon loriciforme (Rosenst) AR Sm combnov equiv Polypodium loriciforme Rosenst RepertSpec Nov Regni Veg 22 17 1925 ndash DistributionCosta Rica to Peru

Serpocaulon maritimum (Hieron) AR Sm comb novequiv Polypodium maritimum Hieron Bot Jahrb Syst34 527 1904 ndash Distribution Nicaragua to Peru

Serpocaulon meniscifolium (Langsd amp Fisch) ARSm comb nov equiv Polypodium meniscifoliumLangsd amp Fisch Pl Voy Russes Monde 11 pl 111810 ndash Distribution S Brazil

Serpocaulon mexiae (Copel) AR Sm comb nov equivPolypodium mexiae Copel Univ Calif Publ Bot17 233 1941 ndash Distribution Minas Gerais Brazil

Serpocaulon nanegalense (Sodiro) AR Sm combnov equiv Polypodium nanegalense Sodiro RecensCrypt vasc quit 65 1883 ndash DistributionColombia Ecuador

Serpocaulon panorense (C Chr) AR Sm comb novequiv Polypodium panorense C Chr Dansk Bot Ark6(3) 97 1929 ndash Distribution Guyana (vide Hensen1990) Brazil

Serpocaulon patentissimum (Mett ex Kuhn) AR Smcomb nov equiv Polypodium patentissimum Mett exKuhn Linnaea 36 134 1869 ndash DistributionColombia Ecuador

Serpocaulon polystichum (Link) AR Sm comb novequiv Polypodium polystichum Link Hort Reg BotBerol 2 101 1833 ndash Distribution Ecuador toBolivia


928

Serpocaulon ptilorhizon (H Christ) AR Sm combnov equiv Polypodium ptilorhizon H Christ Bull HerbBoissier seacuter 2 5 6 1905 ndash Distribution Nicaraguato Bolivia

Serpocaulon richardii (Klotzsch) AR Sm comb novequiv Polypodium richardii Klotzsch Linnaea 20 3941847 ndash Distribution Guianas Venezuela Colombiato Bolivia Brazil

Serpocaulon semipinnatifidum (Feacutee) AR Sm combnov equiv Goniophlebium semipinnatifidum Feacutee MeacutemFoug 5 256 1852 ndash Distribution ColombiaVenezuela Peru

Serpocaulon sessilifolium (Desv) AR Sm comb novequiv Polypodium sessilifolium Desv Meacutem Soc LinnParis 6 238 1827 ndash Distribution Venezuela andColombia to Bolivia

Serpocaulon silvulae (M Kessler amp AR Sm) ARSm comb nov equiv Polypodium silvulae M Kessleramp AR Sm Candollea 60 280 2005 ndashDistribution Bolivia

Serpocaulon subandinum (Sodiro) AR Sm combnov equiv Polypodium subandinum Sodiro Crypt vascquit 348 1893 ndash Distribution Ecuador and Peru

Serpocaulon triseriale (Sw) AR Sm comb nov (Fig3F 4LndashO) equiv Polypodium triseriale Sw J Bot(Schrader) 1800(2) 126 1801 ndash Distribution SMexico to Bolivia Guianas Brazil ParaguayAntilles Trinidad

Serpocaulon vacillans (Link) AR Sm comb nov equivPolypodium vacillans Link Hort berol 2 97 1833ndash Distribution Bolivia S Brazil Paraguay

Serpocaulon wagneri (Mett) AR Sm comb nov equivPolypodium wagneri Mett Ann Sci Nat Bot seacuter5 2 255 1864 ndash Distribution Costa Rica toVenezuela and Ecuador

Excluded speciesPolypodium kunzeanum C Chr ndash Cited by Tryon and

Stolze (1993) as a synonym of P sessilifolium Desvwhich we believe is not closely related Polypodium kun-zeanum (Costa Rica to Peru and Brazil) has much lessregular venation sometimes with two included and them-selves anastomosing veins within an areole cordate-based non-adnate pinnae (the basalmost with lobes over-lapping the rachis) and non-clathrate rhizome scales

ACKNOWLEDGEMENTSWe thank Charles Alford Ivan Jimenez Robbin Moran

Michael Kessler and staff of The University of CaliforniaBotanical Garden for providing samples of Serpocaulon Thanksalso to Daniel Tejero-Diacuteez and to an anonymous reviewer forreading and commenting on the manuscript This research was

supported by a collaborative grant from the National ScienceFoundation to ARS (DEB-9807053) CHH (DEB-9807054) andTAR (DEB-9726607) as well as support to HS from the GermanScience Foundation (DFG SCHN 7582-1) (under theSchwerpunkt Programm SPP 1127 ldquoRadiationsmdashOrigin ofBiological Diversityrdquo

LITERATURE CITEDDavidse G Sousa M amp Knapp S (gen eds) 1995 Flora

Mesoamericana Vol 1 Psilotaceae a Salviniaceae R CMoran and R Riba (Pteridophyte eds) Univ NacionalAutoacutenoma de Meacutexico Ciudad Universitaria

de la Sota E R 1966 Revisioacuten de las especies americanas delgrupo Polypodium squamatum L Revista Mus La PlataSecc Bot 10 69ndash186

Flora of North America Editorial Committee [FNA EdComm] 1993 Flora of North America North of MexicoVol 2 Pteridophytes and Gymnosperms Oxford UnivPress Oxford

Goacutemez L D 1977 Contribuciones a la pteridologia cen-troamericana 2 Novitates Brenesia 1011 115ndash119

Goacutemez L D 1985 Microphlebodium un nuevo nombregenerico (Polypodiaceae) Phytologia 59 58

Haufler C H Grammer W A Hennipman E Ranker TA Smith A R amp Schneider H 2003 Systematics ofthe ant-fern genus Lecanopteris (Polypodiaceae) testingphylogenetic hypothesis with DNA sequences Syst Bot28 217ndash227

Haufler C H amp Ranker T A 1995 [1996] RbcL sequencesprovide phylogenetic insights among sister species of thefern genus Polypodium Amer Fern J 85 361ndash374

Haufler C H Soltis D E amp Soltis P S 1995 Phylogenyof the Polypodium vulgare complex insights from chloro-plast DNA restriction site data Syst Bot 20 110ndash119

Hensen R V 1990 Revision of the Polypodium loriceumcomplex (Filicales Polypodiaceae) Nova Hedwigia 50279ndash336

Huelsenbeck J P amp Ronquist F 2001 MrBayes Bayesianinference of phylogenetic trees Bioinformatics 17754ndash755

Janssen T amp Schneider H 2005 Exploring the evolution ofhumus collecting leaves in drynarioid ferns(Polypodiaceae Polypodiidae) Pl Syst Evol 252175ndash197

Kessler M amp Smith A R 2005 Seven new species 13 newcombinations and one new name of Polypodiaceae fromBolivia Candollea 60 271ndash288

Kubitzki K (ed) 1990 The Families and Genera of VascularPlants Vol 1 Pteridophytes and Gymnosperms SpringerBerlin

Lellinger D B 1989 The ferns and fern-allies of Costa RicaPanama and the Chocoacute (Part 1 Psilotaceae throughDicksoniaceae) Pteridologia 2A 1ndash364

Lellinger D B 1993 Application of the name Goniophlebiumand a new subgeneric name in Polypodium Amer Fern J83 37ndash38

Maddison D R amp Maddison E P 2000 MacClade 40Sinauer Associates Sunderland


929

Maxon W R 1903 A study of certain Mexican andGuatemalan species of Polypodium Contr US NatlHerb 8 271ndash280

Maxon W R 1916a Studies of tropical American ferns No6 Polypodium furfuraceum and related species ContrUS Natl Herb 17 557ndash579

Maxon W R 1916b Studies of tropical American ferns No6 Polypodium squamatum and its allies Contr US NatlHerb 17 579ndash596

Mickel J T amp Smith A R 2004 The pteridophytes ofMexico Mem New York Bot Gard 88 indashxxiv 1ndash1055

Posada D amp Crandall K A 1998 Modeltest testing themodel of DNA substitution Bioinformatics 14 817ndash818

Roumldl-Linder G 1990 A monograph of the fern genusGoniophlebium (Polypodiaceae) Blumea 34 277ndash423

Schneider H Smith A R Cranfill R Haufler C HRanker T A amp Hildebrand T 2002 Gymnogrammitisdareiformis is a polygrammoid fern (Polypodiaceae)mdashresolving an apparent conflict between morphological andmolecular data Pl Syst Evol 234 121ndash136

Schneider H Janssen T Hovenkamp P Smith A RCranfill R Haufler C H amp Ranker T A 2004aPhylogenetic relationships of the enigmatic Malesian fernThylacopteris (Polypodiaceae Polypodiidae) Int J PlSci 165 1077ndash1087

Schneider H Smith A R Cranfill R Hildebrand T EHaufler C H amp Ranker T A 2004b Unraveling thephylogeny of polygrammoid ferns (Polypodiaceae ampGrammitidaceae) exploring aspects of the diversificationof epiphytic plants Molec Phylog Evol 31 1041ndash1063

Smith A R amp Mickel J T 1977 Chromosome counts forMexican ferns Brittonia 29 391ndash398

Sorsa V 1966 Chromosome studies in the PolypodiaceaeAmer Fern J 56 113ndash119

Swofford D L 2000 PAUP Phylogenetic Analysis usingParsimony ( and other methods) version 4 SinauerAssociates Sunderland

Tejero-Diacuteez J D amp Pacheco L 2004 Taxa nuevos nomen-clatura redefinicioacuten y distribucioacuten de las especies rela-cionadas con Polypodium colpodes Kunze (Polypodia-ceae Pteridophyta) Acta Bot Mex 67 75ndash115

Tryon A F amp Lugardon B 1991 Spores of the Pteridophy-ta Springer-Verlag New York

Tryon R M amp Stolze R G 1993 Pteridophyta of Peru PartV 18 Aspleniaceaendash21 Polypodiaceae Fieldiana Botns 32 1ndash190

Tryon R M amp Tryon A F 1982 Ferns and Allied Plantswith Special Reference to Tropical America SpringerBerlin

Walker T G 1966 A cytotaxonomic survey of the pterido-phytes of Jamaica Trans Roy Soc Edinburgh 66 169ndash237

Walker T G 1985 Cytotaxonomic studies of the ferns ofTrinidad 2 The cytology and taxonomic implicationsBull Brit Mus (Nat Hist) Bot 13 149ndash249

Weatherby C A 1939 The group of Polypodium polypodi-oides Contr Gray Herb 124 22ndash35

Weatherby C A 1947 Polypodium lepidopteris and its rela-tives in Brazil Contr Gray Herb 16576ndash82

Wendt T 1980 Notes on some Pleopeltis and Polypodiumspecies of the Chihuahuan Desert Region Amer Fern J70 5ndash11

Windham M D 1993 New taxa and nomenclatural changesin the North American fern flora Contr Univ MichiganHerb 19 31ndash61


930

Appendix 1 Voucher information and GenBank accession numbers given for samples of Serpocaulon used in thisstudySpecies origin voucher GenBank accession no (rbcL rps4 trnLF)S appressum Bolivia Jimenez 862 (LPB) DQ151905 DQ151930 DQ151957 S attenuatum Nicaragua Moreno 10406 (UC)DQ151912 DQ151937 DQ151964 S crystalloneuron Bolivia Jimenez 1109 (LPB) DQ151906 DQ151931 DQ151958 S crystal-loneuron Bolivia Jimenez 1176 (LPB) DQ151907 DQ151932 DQ151959 S dissimile Ecuador Smith 2852 (UC) DQ151908DQ151933 DQ151960 S eleutherophlebium Ecuador Smith 2877 (UC) AY 362601 AY362673 DQ151983 S fraxinifolium BoliviaJimenez 696 (UC) DQ151909 DQ151934 DQ151961 S giganteum Bolivia Jimenez 1035 (UC) DQ151910 DQ151935 DQ151962S gilliesii Bolivia Ehrich 314 (UC) DQ151911 DQ151936 DQ151937 S intricatum Bolivia Jimenez 694 (UC) DQ151919DQ151944 DQ151967 S latipes (intricatum) Bolivia Jimenez 1324 (LPB) DQ151915 DQ151940 DQ151971 S lasiopus BoliviaLewis 35186 (UC) DQ151913 DQ151938 DQ151965 S latipes Bolivia Bach 1305 (LPB) DQ151914 DQ151939 DQ151966 Slatissimum Bolivia Jimenez 813 (UC) DQ151916 DQ151941 DQ151968 S levigatum Bolivia Jimenez 756 (UC) DQ151917DQ151942 DQ151969 S maritimum cult Alford Smith sn (UC) DQ151918 DQ151943 DQ151970 S patentissimum EcuadorSmith 2829 (UC) AY362602 AY362674 - S ptilorhizon Bolivia Jimenez 1102 (UC) DQ151920 DQ151945 DQ151972 S richardiiBolivia Jimenez 1986 (UC) DQ227291 DQ151929 DQ151956 S richardii Ecuador Moran 6258 (NY) AY362593 AY362666 - Ssessilifolium Bolivia Jimenez 755 (LPB) DQ151923 DQ151948 DQ151975 S sessilifolium Bolivia Jimenez 806 (LPB) -DQ151951 DQ151978 S sessilifolium Bolivia Jimenez 976 (UC) DQ151924 DQ151949 DQ151976 S sessilifolium BoliviaJimenez 1034 (LPB) - DQ151950 DQ151977 S subandinum Peru Hutchison amp Wright 5872 (UC) DQ151922 DQ151947DQ151974 S subandinum Ecuador Moran 6858 (NY) DQ151921 DQ151946 DQ151973 S silvulae Bolivia Jimenez 1093 (LPB)DQ151925 DQ151952 DQ151979 S triseriale Bolivia Jimenez 1994 (UC) DQ151926 DQ151953 DQ151980 S vacillans BrazilMexia 5856a (UC) DQ151927 DQ151954 DQ151981 S wagneri Venezuela Smith 1171 (UC) DQ151928` DQ151955 DQ151982

Smith amp al bull Serpocaulon a new fern genus55 (4) bull November 2006 Electronic supplement

1

Appendix 2 Synonyms referable to species of Serpocaulon (basionyms only) Names included herein are largely on thebasis of descriptions (original and other) and circumscriptions adopted by Lellinger (1989) Hensen (1990) and a fewother sources as well as provenance of the type and any original illustrationsGoniophlebium acuminatum Feacutee Meacutem Foug 11 68 pl 19 1866 = Serpocaulon sessilifolium (Desv) AR Sm (fide Hensen 1990)Goniophlebium arcuatum Feacutee Meacutem Foug 5 255 1852 = Serpocaulon loriceum (L) AR Sm (fide Hensen 1990)Goniophlebium chondrocheilon Feacutee Meacutem Foug 8 92 1857 = Serpocaulon eleutherophlebium (Feacutee) AR Sm (fide Hensen 1990)Goniophlebium deflexum T Moore amp Houlst Gard Mag Bot 3 61 1851 = Serpocaulon fraxinifolium (Jacq) AR Sm (fide Lellinger

1989)Goniophlebium demissum Feacutee Crypt Vasc Breacutesil 2 63 1872 = Serpocaulon catharinae (Langsd amp Fisch) AR Sm (fide Hensen 1990)Goniophlebium elatum Feacutee Meacutem Foug 5 256 1852 = Serpocaulon dissimile (L) AR Sm (fide Hensen 1990)Goniophlebium excelsior Feacutee Crypt Vasc Breacutesil 1 244 pl 77 1869 = Serpocaulon meniscifolium (Langsd amp Fisch) AR SmGoniophlebium gauthieri Feacutee Crypt Vasc Breacutesil 1 112 t 34 f 3 1869 = Serpocaulon meniscifolium (Langsd amp Fisch) AR Sm (fide

Hensen 1990)Goniophlebium grammitoides Feacutee Crypt Vasc Breacutesil 1 110 pl 34 1869 = Serpocaulon vacillans (Link) AR SmGoniophlebium haenkei C Presl Tent pterid 186 pl 7 1836 = Serpocaulon triseriale (Sw) AR Sm (fide Hensen 1990)Goniophlebium intermedium Feacutee Meacutem Foug 11 69 t 18 f 3 1866 = Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989)Goniophlebium invertens Feacutee Meacutem Foug 9 25 1857 = Serpocaulon falcaria (Kunze) AR Sm (fide Hensen 1990)Goniophlebium pectinans Feacutee Crypt vasc Breacutesil 1 109 pl 34 1869 = Serpocaulon catharinae (Langsd amp Fisch) AR Sm (fide Hensen

1990)Goniophlebium pectinatum J Sm in Seemann Bot Voy Herald 230 1854 (nom illeg non Goniophlebium pectinatum (L) J Sm 1841

which is Pecluma pectinata (L) M G Price) = Serpocaulon wagneri (Mett) AR Sm (fide Lellinger 1989 Hensen 1990)Goniophlebium pictum Feacutee Crypt Vasc Breacutesil 1 244 1869 equiv Polypodium pictum (Feacutee) Brade Arq Inst Biol Veg 1 228 1935 non P

pictum (T Moore) Baker equiv Polypodium limbatum Brade Arq Jard Bot Rio de Janeiro 11 30 1951 = Serpocaulon catharinae (Langsdamp Fisch) AR Sm (fide Hensen 1990)

Polypodium adnatum Kunze ex Klotzsch var glaucescens Mille Revista Col Nac Vicente Rocafuerte 9 208 1927 = Serpocaulon gigan-teum (Desv) AR Sm (fide Lellinger 1989)

Polypodium albopunctatum Raddi Opusc Sci 3 287 1819 = Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989 Hensen 1990)Polypodium andinum H Karst Fl Columb 1 171 pl 85 1861 (non Hook 1860) = Serpocaulon sessilifolium (Desv) AR Sm (fide Lel-

linger 1989 Hensen 1990)Polypodium argyrolepis Sodiro Crypt vasc quit 355 1893 = Serpocaulon lasiopus (Klotzsch) AR Sm (fide Hensen 1990)Polypodium articulatum Desv Meacutem Soc Linn Paris 6 236 1827 (non P articulatum (Sw) Juss ex Poir 1804 which is Oleandra artic-

ulata (Sw) C Presl) = Serpocaulon caceresii (Sodiro) AR Sm (fide Hensen 1990)Polypodium beyerianum Rosenst Repert Spec Nov Regni Veg 22 17 1925 = Serpocaulon loriceum (L) AR Sm (fide Lellinger 1989

but for a different opinion see Hensen 1990 who placed it in synonymy under Polypodium maritimum Hieron)Polypodium brasiliense Poir in Lam Encycl 5 525 1804 = Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989 Hensen 1990)

= Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989)Polypodium brasiliense Poir var pleiosorum Rosenst ex Hassl Treb Inst Bot Farm Buenos Aires 45 71 1928 = Serpocaulon triseriale

(Sw) AR Sm (vide Lellinger 1989)Polypodium campylopodum Klotzsch Linnaea 20 395 1847 = Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989 Hensen 1990)Polypodium catharinae Langsd amp Fisch forma aurita Rosenst Hedwigia 46 142 1906 = Serpocaulon catharinae (Langsd amp Fisch) AR

SmPolypodium catharinae Langsd amp Fisch forma bipinnatifida Rosenst Hedwigia 46 142 1906 = Serpocaulon catharinae (Langsd amp

Fisch) AR SmPolypodium chartaceum Baker J Bot 15 166 1877 = Serpocaulon loriceum (L) AR Sm sl (fide Lellinger 1989)Polypodium chnoodes Spreng Neue Entd 3 6 1822 = Serpocaulon dissimile (L) AR Sm (fide Lellinger 1989 Hensen 1990)Polypodium chnoodes Spreng var minus H Christ Bull Herb Boissier seacuter 2 6 49 1906 = Serpocaulon dissimile (L) AR Sm (fide

Lellinger 1989)Polypodium costaricense H Christ Bull Herb Boissier 4 660 1896 = Serpocaulon wagneri (Mett) AR Sm (fide Lellinger 1989 Hensen

1990)Polypodium cymatodes Kunze Linnaea 23 277 317 1850 = Serpocaulon fraxinifolium (Jacq) AR Sm (fide Lellinger 1989)Polypodium deflexum Houlst amp T Moore Gard Mag Bot 3 60 1851 = Serpocaulon fraxinifolium (Jacq) AR Sm (fide Hensen 1990)Polypodium distans Raddi Pl Bras Nov Gen 1 21 pl 31 1825 (non Kaulf 1824) = Serpocaulon fraxinifolium (Jacq) AR Sm (fide

Lellinger 1989 Hensen 1990)Polypodium elatius Schrad Goumltt Gel Anz 1824 868 1824 = Serpocaulon meniscifolium (Langsd amp Fisch) AR Sm (fide Lellinger

1989 Hensen 1990)Polypodium elegans Cav ex Sw Syn fil 35 1806 (non Poir 1804) = Serpocaulon loriceum (L) AR Sm sl (fide Lellinger 1989)Polypodium fluminense Vell Arq Mus Nac Rio de Janeiro 5 447 1881 = Serpocaulon fraxinifolium (Jacq) AR Sm (fide Hensen 1990)Polypodium fraxinifolium Jacq var fraxinellum H Christ Bull Herb Boissier seacuter 2 5 5 1905 = Serpocaulon fraxinifolium (Jacq) AR

Sm (fide Lellinger 1989)Polypodium fraxinifolium Jacq subsp articulatum H Christ Bull Herb Boissier seacuter 2 6 49 1906 = Serpocaulon giganteum (Desv) AR

Sm (fide Lellinger 1989)Polypodium fraxinifolium Jacq subsp luridum H Christ Bull Herb Boissier seacuter 2 6 48 1906 = Serpocaulon giganteum (Desv) AR Sm

(fide Lellinger 1989)Polypodium gladiatum Kunze Linnaea 9 45 1834 (non Vell 1827) equiv Polypodium ampliatum Maxon Contr US Natl Herb 10 492 1908

= Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989 Hensen 1990)Polypodium glandulosissimum Brade Arq Inst Biol Veg 1 230 1935 = Serpocaulon catharinae (Langsd amp Fisch) AR Sm (fide

Hensen 1990)

Smith amp al bull Serpocaulon a new fern genus 55 (4) bull November 2006 Electronic supplement

2

Polypodium glaucophyllum Kunze ex Klotzsch Linnaea 20 393 1847 = Serpocaulon levigatum (Cav) AR Sm (fide Lellinger 1989Hensen 1990)

Polypodium glaucum Raddi Pl Bras Nov Gen 20 pl 29 1825 (non Thunb 1784) equiv Chrysopteris raddiana Feacutee Crypt vasc Breacutesil 1 1201869 = Serpocaulon catharinae (Langsd amp Fisch) AR Sm (fide Hensen 1990)

Polypodium harpeodes Link Hort berol 2 97 1833 = Serpocaulon vacillans (Link) AR Sm (fide Hensen 1990 who first synonymizedP harpeodes under P vacillans)

Polyhpodium hassleri H Christ in Hassler Bull Herb Boissier seacuter 2 7 923 1907 = Serpocaulon vacillans (Link) AR SmPolypodium hirsutulum Raddi Pl bras nov gen 1 21 pl 29 1825 = Serpocaulon meniscifolium (Langsd amp Fisch) AR Sm (fide Hensen

1990)Polypodium kuhnii E Fourn Bull Soc Bot France 19 251 1872 = Serpocaulon attenuatum (Humb amp Bonpl ex Willd) AR SmPolypodium laetum Raddi Opusc Sci 3 287 1819 (non Salisb 1796) = Serpocaulon latipes (Langsd amp Fisch) AR Sm (fide Hensen

1990)Polypodium longifolium C Presl Delic prag 1 167 1822 (non Cav 1802) equiv Polypodium preslianum Spreng Syst Veg ed 4 index

1828 = Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989 Hensen 1990)Polypodium loriceum L forma duplosorum Domin Rozpr Kraacutel Ceskeacute Spolecn Nauk Tr Mat-Prir 2[Pterid Dominica] 130 1929 as

ldquoduplisorardquo = Serpocaulon loriceum (L) AR Sm sl (fide Lellinger 1989)Polypodium loriceum L var heterolepis Sodiro Anales Univ Centr Ecuador 11(72) 28 (Crypt vasc quit 352) 1894 = Serpocaulon

loriceum (L) AR Sm sl (fide Lellinger 1989)Polypodium loriceum L var obscurum Rosenst Repert Spec Nov Regni Veg 6 308 1909 as ldquoobscurardquo = Serpocaulon loriceum (L) AR

Sm sl (fide Lellinger 1989)Polypodium loriceum L var oligomerum H Christ Bull Soc Roy Bot Belgique 35 Meacutem 221 1896 = Serpocaulon loriceum (L) AR

Sm sl (fide Lellinger 1989)Polypodium loriceum L var squamuligera Rosenst Repert Spec Nov Regni Veg 6 308 1909 = Serpocaulon loriceum (L) AR Sm sl

(fide Lellinger 1989)Polypodium lucens Schrad Goumltt Gel Anz 1824 868 1824 = Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989 Hensen 1990)Polypodium martii Colla Herb pedem 6 230 1836 = Serpocaulon vacillans (Link) AR Sm (fide Hensen 1990)Polypodium mindense Sodiro Crypt vasc quit 348 1893 = Serpocaulon eleutherophlebium (Feacutee) AR Sm (fide Hensen 1990)Polypodium mosenii C Chr Bot Tiddskr 25 78 1902 = Serpocaulon meniscifolium (Langsd amp Fisch) AR Sm (fide Hensen 1990)Polypodium mutabile Kunze Linnaea 9 46 1834 = Serpocaulon fraxinifolium (Jacq) AR Sm (fide Lellinger 1989 Hensen 1990)Polypodium neriifolium Schkuhr 24 Kl Linn Pfl-Syst 1 14 t 15 1804 = Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989

Hensen 1990)Polypodium neriifolium Schkuhr var acuminatissimum Kuntze Revis gen pl 2 819 1891 = Serpocaulon triseriale (Sw) AR Sm (fide

Lellinger 1989 holotype NY)Polypodium neriifolium Schkuhr var heterophyllum Kuntze Revis gen pl 2 819 1891 = Serpocaulon triseriale (Sw) AR Sm (fide Lel-

linger 1989 holotype NY)Polypodium ornatum Klotzsch Linnaea 20 396 1847 = Serpocaulon fraxinifolium (Jacq) AR Sm (fide Hensen 1990)Polypodium palma Vell Arq Mus Nac Rio de Janeiro 5 447 1881 = Serpocaulon meniscifolium (Langsd amp Fisch) AR Sm (fide

Hensen 1990)Polypodium preslianum var immersum Rosenst Repert Spec Nov Regni Veg 6 314 1909 (as ldquoimmersardquo) = Serpocaulon triseriale (Sw)

AR Sm (fide Lellinger 1989)Polypodium pseudofraternum AC Sm Bull Torrey Bot Club 58 307 1931 = Serpocaulon sessilifolium (Desv) AR Sm (fide Lellinger

1989 Hensen 1990 holotype NY)Polypodium punctulatum Hook Icon pl pl 720 1845 (non Poir 1804) = Serpocaulon subandinum (Sodiro) AR Sm (fide Hensen 1990)Polypodium pycnosorum Link Fil spec 128 1841 = Serpocaulon triseriale (Sw) AR Sm (fide Hensen 1990)Polypodium ramosum Loddiges ex Link Hort berol 2 97 1833 = Serpocaulon loriceum (L) AR Sm sl (fide Lellinger 1989)Polypodium remotum Baker Ann Bot (Oxford) 5 470 1891 (non Desv 1827) equiv Polypodium uniseriale C Chr Index fil 527 1906 =

Serpocaulon sessilifolium (Desv) AR Sm (fide Lellinger 1989 Hensen 1990)Polypodium retrofractum Desv Meacutem Soc Linn Paris 6 237 1827 = Serpocaulon dissimile (L) AR Sm (fide Lellinger 1989 Hensen

1990)Polypodium rhizocaulon Willd Sp pl ed 4 5 196 1810 = Serpocaulon fraxinifolium (Jacq) AR Sm (fide Lellinger 1989)Polypodium rimbachii Sodiro Crypt vasc quit 356 1893 = Serpocaulon sessilifolium (Desv) AR Sm (fide Hensen 1990)Polypodium rupincolum Brade Arq Inst Biol Veg 1 230 1935 = Serpocaulon catharinae (Langsd amp Fisch) AR Sm (fide Hensen

1990)Polypodium scutulatum Sodiro Anales Univ Centr Ecuador 22 169 1908 = Serpocaulon fraxinifolium (Jacq) AR Sm (fide Hensen

1990)Polypodium sessilifolium Desv var papallactense Mille Revista Col Nac Vicente Rocafuerte 9 211 1927 = Serpocaulon sessilifolium

(Desv) AR Sm (fide Lellinger 1989)Polypodium subalatum Klotzsch Linnaea 20 394 1847 = Serpocaulon triseriale (Sw) AR Sm (fide Hensen 1990)Polypodium subviride Lellinger Proc Biol Soc Wash 98 389 f 17 1985 = Serpocaulon loriciforme (Rosenst) AR SmPolypodium surucuchense Hook Icon pl 1 t 69 1837 = Serpocaulon sessilifolium (Desv) AR Sm (fide Hensen 1990)Polypodium vagans Mett Ann Sci Nat Bot seacuter 5 2 255 1864 = Serpocaulon eleutherophlebium (Feacutee) AR Sm (fide Hensen 1990)Polypodium variolatum Willd Sp pl ed 4 5 192 1810 = Serpocaulon triseriale (Sw) AR Sm (fide Lellinger 1989 Hensen 1990)Polypodium wiesbaueri Sodiro Recens Crypt vasc quit 65 1883 = Serpocaulon eleutherophlebium (Feacutee) AR Sm (fide Hensen 1990)Polypodium xiphophoron Kunze ex Mett Abh Senckenberg Naturf Ges 2 73 1856 = Serpocaulon triseriale (Sw) AR Sm (fide Hen-

sen 1990)

Appendix 2 (continued)

(Maxon 1916a 21 spp Groups 4 and 6 parts of Tryonamp Tryon 1982) 4) P polypodioides (Maxon 1916bWeatherby 1939 Group 4 part of Tryon amp Tryon1982) 5) P squamatum and relatives (Maxon 1916bWeatherby 1947 revised by de la Sota 1966 22 sppGroup 5 part of Tryon amp Tryon 1982) 6) the P thys-sanolepis group (Wendt 1980 Group 4 part of Tryon ampTryon 1982) and 7) P loriceum P fraxinifolium andallies an entirely neotropical group of about 40 specieswith regularly anastomosing (goniophlebioid) veins andoften long-creeping cordlike rhizomes bearing partiallyor entirely clathrate scales (most of Group 2 of Tryon ampTryon 1982 species taxonomy 23 spp revised byHensen 1990) Because nearly all of the major sub-groups of Polypodium sl occur in Mexico and CentralAmerica and these areas are a center of diversity for sev-eral of the subgroups many species have been treated infloristic contexts by Lellinger (1989) Moran (in Davidseamp al 1995) and by Mickel amp Smith (2004) Howeverthe species allied to P loriceum and P fraxinifolium arepredominantly South American and are less well studiedexcept for work by Hensen (1990) which was followedin the main in the floristic account for Peru by Tryon ampStolze (1993)

Existing evidence (Schneider amp al 2004b) supportsthe placement of species in 3) through 6) as delineatedabove in Pleopeltis All of these groups contain speciesthat bear scales on the abaxial lamina A few scaly-blad-ed species of Polypodium have already been transferredto Pleopeltis (Windham 1993) as precursor to theirtreatment in the Flora of North America North of Mexico(Andrews amp Windham in FNA Ed Comm 1993) how-ever most scaly species remain in Polypodium to betransferred to Pleopeltis elsewhere (Smith amp al inprep) Species now placed in the segregate generaDicranoglossum Neurodium Microphlebodium andPseudocolysis also appear to be closely allied to and per-haps inseparable from Pleopeltis sl In the present paperwe address the species mentioned under 7) above andthese lack a name at generic rank This new genus wehere name Serpocaulon alluding to the long-creepingsnakelike rhizomes found in nearly all species

Many of the species of Serpocaulon were initiallydescribed in Goniophlebium (Blume) C Presl (seeAppendix 2 electronic supplement) a genus of about 25species occurring from India throughout southeast andeastern Asia the Himalayan region and Malesia toAustralasia Melanesia Samoa and Micronesia (Roumldl-Linder 1990) The name Goniophlebium is derived fromthe Greek gonia meaning angle and phleps vein refer-ring to the characteristic venation pattern formed by reg-ularly anastomosing veins that often form a succession ofchevron-shaped areoles between the pinna midribs andmargins Although the paleotropical species of Gonio-

phlebium and the neotropical species of Serpocaulonhave similar venation and both genera bear clathrate rhi-zome scales the similarities are convergent as indicatedby their positions in phylogenetic trees based on molec-ular data (see Schneider amp al 2004b) Goniophlebiumss falls within an almost exclusively monophyletic OldWorld clade (as far as known the lone exception is a sin-gle disjunct species of Platycerium in the AndesSchneider amp al 2004b) whereas Serpocaulon nests in apredominantly New World clade (with a few African out-liers) containing the genera Microgramma Campyloneu-rum Pleopeltis Pecluma and Polypodium ss as wellas the pantropical grammitid clade Roumldl-Linder (1990)and Lellinger (1993) also specifically excluded all NewWorld species of Polypodium from Goniophlebium onthe basis of several morphological characters eg theabsence of black sclerenchyma strands in the rhizomespeltate-based rhizome scales the absence of hairy orscaly paraphyses in the sori and the folded perine aroundthe spores

In this paper we restrict the application ofPolypodium L (Sp pl 2 1082 1753) with lectotype Pvulgare L (chosen by John Smith Hist fil 88 1875) toa group of predominantly temperate (mainly north-tem-perate) species plus a less widespread tropical and sub-tropical group of Mexican-Mesoamerican speciesPolypodium as defined in this narrow sense then com-prises about 15 species in temperate regions of AmericaEurope Asia and Africa (Haufler amp al 1995 Haufler ampRanker 1995) and perhaps another 15 species in Mexicoand Mesoamerica The species have deeply pinnatifid orweakly 1-pinnate blades concolorous or sometimesslightly bicolorous orange-tan to brown non-clathraterhizome scales free or weakly anastomosing venation(each areole with a single free included veinlet) soriterminal on ends of free included veinlets channeledstipes filamentous or branched paraphyses and morefrequently branched rhizomes

Within Serpocaulon eight (of about 40 total) specieshave been included previously (as Polypodium) in a phy-logenetic study using the chloroplast DNA regions rbcLrps4 and rps4-trnS intergenic spacer (Schneider amp al2004b) Polypodium adnatum Kunze ex Klotzsch[voucher re-identified herein as S richardii (Klotzsch)Domin] P fraxinifolium Jacq P funckii Mett (voucherre-identified as S ptilorhizon (H Christ) A R Sm) butnot used in the current study) P levigatum Cav Ppatentissimum Mett ex Kuhn P ptilorhizon H Christ Ptriseriale Sw and P wiesbaueri Sodiro [= Peleutherophlebium (Feacutee) Mett in this paper] In a maxi-mum likelihood analysis (Schneider amp al 2004b) thisclade is monophyletic with clade confidence values of100 (Bayesian posterior probability PP) and non-para-metric bootstrap support of 99 In the Schneider amp al


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serpocaulon (polypodiaceae), a new genus segregated from...

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