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Molecular phylogenetics and generic taxonomy of Blechnaceae ferns

August 2014
Taxon 63(4)

DOI:10.12705/634.13

Project: Flora of New Zealand ferns

Authors:

Leon Perrie

Museum of New Zealand, Te Papa Tongarewa

Lara D Shepherd

Museum of New Zealand, Te Papa Tongarewa

Daniel Ohlsen

Royal Botanic Gardens Victoria

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The fern family Blechnaceae is cosmopolitan; however, the vast majority of species are placed in Blechnum, which occurs predominantly in the Southern Hemisphere. There are two areas that are particularly species-rich: the south-west Pacific (including Australasia), and Central and South America. Using chloroplast DNA sequences, we report the first comprehensive phylogenetic analysis of the Blechnaceae, including all genera widely recognised in recent treatments, and over half of the species. There is strong support for several major clades, which we characterise morphologically and geographically, and some of their interrelationships. Blechnum is confirmed as polyphyletic. Blechnum indicum and B. serrulatum are more closely related to Salpichlaena and Stenochlaena, and are segregated as a new genus, Telmatoblechnum. Alternative generic circumscriptions are discussed for the remainder of Blechnum. In the absence of morphological characters to diagnose the clades within core Blechnum, and for the sake of taxonomic stability, we advocate a broad circumscription for Blechnum. Brainea and Sadleria are retained as their relationships are not well resolved, but Doodia and Pteridoblechnum are clearly nested within the core of Blechnum and we provide four new names in Blechnum. Additionally, given the focus of our sampling, we discuss the biogeography of the south-west Pacific, where immigration has been more important in community assembly than what might be superficially inferred from patterns of endemism (with ca. 60% of species endemic to individual land areas).

Blechnaceae phylogeny based on analyses of chloroplast DNA sequences for the full sample set. Maximum likelihood tree, with maximum parsimony bootstrap, maximum likelihood bootstrap, and Bayesian posterior probability given for major clades and the deeper branches. Other branches are bolded if posterior probability ≥ 0.95 and maximum parsimony and/or maximum likelihood bootstrap support ≥ 80%. Species are labelled according to the taxonomy advocated here, with recent synonyms in brackets. Blechnum is abbreviated in species names, but names of other genera are given in full. Sample origin is indicated where known; otherwise the species' distribution is given in square brackets. Amer. = North, Central, and/or South America.

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Phylogenetic summary of the Blechnaceae, with taxonomic options A-G for generic boundaries as discussed in the text. See Fig. 1 for definition of clades/ lineages, and Table 1 for their characterisation. Support values: maximum parsimony bootstrap support, maximum likelihood bootstrap support, Bayesian posterior probability. The branch uniting the Serrulatum clade and Stenochlaena was not recovered in the maximum parsimony analysis.

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Characterisation of the major Blechnaceae clades. See Fig. 1 for circumscription of the clades.

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

Perrie & al. • Blechnaceae phylogeny and taxonomy

TAX O N



Version of Record (identical to print version).

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



INTRODUCTION

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











            





 

Blechnum      





 









Blechnum



DoodiaDoodia

    



Blechnum

Doodia



         

Woodwardia        

     

      



Molecular phylogenetics and generic taxonomy of Blechnaceae ferns

Leon R. Perrie,1 Ruby K. Wilson,2 Lara D. Shepherd,1,3 Daniel J. Ohlsen,2 Erin L. Batty,2

Patrick J. Brownsey1 & Michael J. Bayly2

 Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand

 School of Botany, The University of Melbourne, Melbourne, Australia

 School of Biological Sciences, Victoria University of Wellington, Wellington, New Zealand

 Leon Perrie, leon.perrie@tepapa.govt.nz

DOI



Abstract

Blechnum







 

 Blechnum     Blechnum indicumB. serrulatum  

SalpichlaenaStenochlaenaTelmatoblechnum

 Blechnum 

BlechnumBlechnum. BraineaSadleria

DoodiaPteridoblechnum 

BlechnumBlechnum



  

Keywords

BlechnumDoodia  Telmatoblechnum

Supplementary Material







TAX O N

Perrie & al. • Blechnaceae phylogeny and taxonomy

Version of Record (identical to print version).

   

era Anchistea Lorinseria 

Woodwardia    Sadle-

ria      

Stenochlaena  



Salpichlaena 

   Pteridoblechnum

    

     Steenisio-

blechnum

 Brainea



Woodwardia      

  



     







 trnL-trnF 



 



trnL-trnFtrnG-trnR

  

   



         



 

     





   



MATERIALS AND METHODS





  Blechnum Brainea Doodia Pteridoblechnum

SteenisioblechnumSadleria SalpichlaenaSteno-

chlaena WoodwardiaAnchistea Lorin-

seria



    Blechnum  Doodia  

  

trnL-trnF 

    rbcL  

      rbcL rps4

rps4-trnS   

   

   





      

 rbcL rps4  rps4-trnS 

   trnL-trnFtrnL  trnL  

  trnL-trnF 

     



   

   Doodia

hindiiD. maxima 

  

  

    

trnL-trnF

       



  

     









  



rbcL 





  







     

   

    

   

rbcLtrnL-trnF  

   



 rps4 rps4-trnS  



  



    



 







  

trnL-trnFrps4-trnS

 







      

 

    

     





Perrie & al. • Blechnaceae phylogeny and taxonomy

TAX O N



Version of Record (identical to print version).

        

 





               

  





rbcLtrnL-trnF 











     







 







      



   

      









  







RESULTS

       







 













       







    

Stenochlaena SalpichlaenaBlechnum

indicum  B. serrulatumBlech-

num  

Stenochlaena  

 Salpichlaena  Stenochlaena





     

Wo o d-

wardia Stenochlaena

SalpichlaenaBlechnum indicum B. serrulatum







   Blechnum 



      

        







  

BlechnumB. occidentale

DoodiaPteri-

doblechnum B. articu-

latum

      











Brainea Sadleria Blechnum

   Blechnum

eburneum









   

    





DISCUSSION







   

          

    

  





   

   Woodwardia   

 

      



TAX O N

Perrie & al. • Blechnaceae phylogeny and taxonomy

Version of Record (identical to print version).

Super-

Stenochlaena

-,28,-

Woodwardia areolata, U.S.A.

Woodwardia virginica, U.S.A.

Woodwardia japonica, Japan

Woodwardia radicans, ? [Europe]

Woodwardia prolifera, ? [Asia]

Salpichlaena volubilis, Costa Rica

Telmatoblechnum indicum (B. indicum), Australia

Telmatoblechnum serrulatum (B. serrulatum), ? [Amer.]

Stenochlaena palustris, Fiji

Stenochlaena palustris, Australia

Stenochlaena tenuifolia, ? [Africa]

Stenochlaena milnei, Philippines

Brainea insignis, ? [Asia]

B. spicant, ? [Europe, Amer.]

B. amabile, Japan

B. hancockii, Taiwan

Sadleria squarrosa, Hawaii

Sadleria cyatheoides, Hawaii

Sadleria soulayetiana, Hawaii

B. eburneum, China B. orientale, Australia

B. orientale, Fiji

B. orientale, New Caledonia

B. orientale, Japan

B. microbasis, ? [Africa]

B. simillimum, ? [Africa]

B. contiguum, New Caledonia

B. contiguum, Lord Howe I. B. kunthianum, ? [Amer.]

B. fragile, Costa Rica

B. ensiforme, ? [Amer.]

B. binervatum, ? [Amer.]

B. nudum, Australia

B. discolor, New Zealand

B. oceanicum, New Caledonia

B. filiforme, New Zealand B. sampaioanum, Brazil

B. opacum, New Caledonia

B. fullagari, Lord Howe I.

B. nigrum, New Zealand

B. fluviatile, Australia

B. fluviatile, New Zealand

B. hirsutum, New Caledonia

B. vulcanicum, Australia

B. vulcanicum, New Zealand

B. pilosum, Fiji

B. punctulatum, ? [Africa]

B. australe, ? [Amer.] B. polypodioides, ? [Amer.]

B. austrobrasilianum,? [Amer.]

B. laevigatum, Argentina

B. glandulosum, ? [Amer.]

B. gracile, ? [Amer.]

B. appendiculatum, ? [Amer.]

B. occidentale, ? [Amer.]

B. mochaenum, Argentina

B. blechnoides, New Zealand

B. durum, New Zealand

B. penna-marina, New Zealand

B. penna-marina, Australia

B. penna-marina, Argentina

B. microphyllum, Argentina

B. aff. chambersii, Fiji

B. chambersii, Australia

B. chambersii, New Zealand

B. norfolkianum, New Zealand

B. membranaceum, New Zealand

B. colensoi, New Zealand

B. melanocaulon, Fiji

B. vieillardii, New Caledonia

B. patersonii subsp. queenslandicum, Australia

B. patersonii subsp. patersonii, Australia

Contiguum

Super-Blechnum

Core-Blechnum

Occidentale

100,100,1.0

94,96,1.0

87,87,1.0

100,100,1.0

-,76,0.99

92,99,1.0

91,97,1.0

97,100,1.0

92,97,1.0

Serrulatum=Telmatoblechnum

Salpichlaena

Stenochlaena

Woodwardia

60,77,1.0

-,26,0.76

-,61,0.95

0.01

Nudum

88,85,1.0

96,100,1.0

-,48,0.82

-,53,0.91

-,38,0.88

Amabile

Sadleria

Onocleopsis hintonii, Mexico

Pentarhizidium intermedium, China

Onoclea sensibilis, ? [Amer., Asia]

Matteuccia struthiopteris, ? [Amer., Asia, Europe]

100,100,1.0

75,79,0.98

Onocleaceae

outgroup

continued next page



Perrie & al. • Blechnaceae phylogeny and taxonomy

TAX O N



Version of Record (identical to print version).

         









 







        



   





  Blechnum



   

    

Woodwardia    

 





B. novae-zelandiae, New Zealand

B. minus, New Zealand

B. howeanum, Lord Howe I.

B. montanum, New Zealand

B. spinulosum (Doodia caudata), Australia

B. brasiliense, [Amer.]

B. cartilagineum, Australia

B. whelanii, Australia

B. gibbum, New Caledonia

B. moorei, New Caledonia

B. obtusatum, New Caledonia

B. vittatum, Fiji

B. parrisii (Doodia australis), New Zealand

B. neohollandicum (Doodia aspera), Australia

B. medium (Doodia media), New Caledonia

B. norfolkense (Doodia kunthiana), Hawaii

B. parrisii (Doodia australis), Australia

B. medium (Doodia media), Australia

B. brackenridgei (Doodia brackenridgei), Fiji

B. dissectum (Doodia dissecta), Australia

B. kermadecense (Doodia milnei), New Zealand

B. doodianum (Doodia heterophylla), Australia

B. zeelandicum (Doodia squarrosa), New Zealand

B. molle (Doodia mollis), New Zealand

B. lineare (Doodia linearis), Australia

B. fraseri, New Zealand B. neglectum (Pteridoblechnum neglectum), Australia

B. reticulatum (Pteridoblechnum acuminatum), Australia

B. diversifolium, New Caledonia

B. lenormandii, New Caledonia

B. schomburgkii, ? [Amer.]

B. magellanicum, ? [Amer.]

B. moritzianum, Argentina

B. tabulare, Argentina B. articulatum, Australia

B. gregsonii, Australia

B. camfieldii, Australia

B. wurunuran, Australia

B. procerum, New Zealand

B. wattsii, Australia

B. ambiguum, Australia

B. schiedeanum, ? [Amer.]

B. cordatum, ? [Amer.]

B. loxense, Peru

B. stipitellatum, Ecuador

B. corbassonii, New Caledonia

B. subcordatum, New Caledonia

B. chilense, ? [Amer.]

B. milnei, Fiji

B. minus, Australia

B. triangularifolium, New Zealand

Super-Blechnum

Core-Blechnum

0.01

100,100,1.0

92,99,1.0

100,100,1.0

98,98,1.0

Gibbum+Doodia

Procerum

Fraseri

Tabulare

-,21,-

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Fig. 1.

                               



  

 Blechnum 

 





TAX O N

Perrie & al. • Blechnaceae phylogeny and taxonomy

Version of Record (identical to print version).



     



Taxonomy of the super-Stenochlaena clade, including rec-

ognition of a n ew genus. —





    

   Blechnum indicum 

B. serrulatum   

StenochlaenaSalpichlaena  Blechnum



      

Blechnum

 Steno chlaenaSalpichlaena  







BlechnumWoodwardia.







Blechnum indicum B. serrulatum  

 Stenochlaena

      

Salpichlaena



   Stenochlaena

Salpichlaena   Blechnum  



  Stenochlaena  Blechnum 





B. indicum B. serrulatum

Steno-

chlaena    



Stenochlaena    





 Blechnum 



Stenochlaena



 B. indicum B. serrulatum

Telmatoblechnum



Taxonomy of the super-Blechnum clade, including syn-

onymisation of Doodia and Pteridoblechnum under Blech-

num. —



    Blechnum   B. indicum

Tabulare

Nudum

Fraseri

Serrulatum

Sadleria

Brainea

Spicant

Orientale

Eburneum

Amabile

Contiguum

Procerum

Occidentale

Gibbum

+Doodia

Woodwardia

Salpichlaena

Stenochlaena

100,100,1.0

-,76,0.99

97,100,1.0

100,100,1.0

94,96,1.0

60,77,1.0

87,87,1.0

91,97,1.0

88,85,1.0

96,100,1.0

92,99,1.0

100,100,1.0

92,99,1.0

100,100,1.0

92,97,1.0

98,98,1.0

B C D E F GA

Taxonomic options

Super-Blechnum

Core-Blechnum

Super-

Stenochlaena

75,79,0.98

Fig. 2.













 

  







 

Stenochlaena







Perrie & al. • Blechnaceae phylogeny and taxonomy

TAX O N



Version of Record (identical to print version).

B. serrulatum     Brainea Doodia

PteridoblechnumSadleria  

BlechnumDoodiaPteridoblechnum

  

          

   











Blechnum    



   







 

     

Blechnum

   



      

     

          

   





                 

 

B. occiden-

tale

  

 

    





  Blechnum 





Table 1.











a







c



 

 





x =

2n =











































x =

n =

2n =







































 



B. inflexum





x =

n =

 























Blechnum











x =

B. occidentale





n =



2n =

























B. occidentale 









B. punctulatum











 







x =

B. articulatum

n =

2n =











B. articulatum

















TAX O N

Perrie & al. • Blechnaceae phylogeny and taxonomy

Version of Record (identical to print version).

 





  











B. columbiense













 





 



















 



x =

2n =

 















 

  

x = 33

2n =

 















 





x = 33

2n =

 

















 



  

x = 34

2n =

 













Brainea 



x = 33

2n =





















Sadleria 



x = 33

n































Telmato-

blechnum









x =

2n = 72

























Stenochlaena 





x = ca. 37

2n =























Salpichlaena 

x = 40

n = 40

























Woodwardia 





x =

2n =

 



















a

nn



c

Table 1.











a







c



 



Perrie & al. • Blechnaceae phylogeny and taxonomy

TAX O N



Version of Record (identical to print version).







   

    

  

Blechnum





       

   Blechnum

   

 

DoodiaPteridoblechnum

Blechnum      

   Doodia  Pteridoblechnum



 Doodia   

 

 Doodia

Blechnum   

 



   Blechnum  

Doodia     



Doodia 

Doodia 



         





    Pteridoblechnum    

   Steenisioblechnum  

Blechnum

fraseri



Blechnum fraseri  

   

 Diploblechnum  

PteridoblechnumPteridoblechnum

    

     

Blechnum

Pteridoblechnum

Blechnum 

           

Blechnum

Pteridoblechnum.

 

Blechnum





   



Blechnum     



Brainea Sadleria



BlechnopsisBlechnum orientaleStruthiopteris

B. spicant   

B. eburneum   

BraineaSadleria   





 B. orientale 

B. finlaysonianum





Blechnum melanopus

Blechnidium 

           

  

BraineaSadleria

   Blechnum  

           

    



    

  Brainea    Sadleria

   

Blechnum  

   Blechnum  

 

BlechnumBrainea   Blechnum insigne

 

Sadleria

Biogeography. —

     

   





 



 

Blech-

num



 Woodwardia  





     

    

   





 

  



           



    



     

     



TAX O N

Perrie & al. • Blechnaceae phylogeny and taxonomy

Version of Record (identical to print version).



      









    



B. chambersiiB. norfolki-

anum B. fluviatile

  B. microphyllum

 B. penna-marina 

B. minus   

& B. novae-zelandiae

B. orientale 

B. procerumB. wattsii 

B. vulcanicum 

Doodia 











   

Woodwardia

Stenochlaena

Brainea   

   Stenochlaena

Woodwardia    

     Stenochlaena Telmato-

blechnumWoodwardia



StenochlaenaTelmatoblechnum    

Salpi-

chlaena Telmatoblechnum

   TelmatoblechnumWoodwardia

   



 StenochlaenaTel -

matoblechnum

  

     

Sadleria



          



   

    







     





          

     

Blechnum corbassoniiB. subcordatum     

B. chauliodontum B. confusum   









B. melanocaulon      

   

B. patersonii





    

Sadleria 





Future work with species complexes. —

 

     



 

 



Blechnum ambiguum 

B. greg soniiB. wattsii

B. minusB. minus 

B. novae-zelandiae     

B. chambersii    Doodia







B. patersonii

B. contiguum

Steno-

chlaena palustris 

TAXONOMIC TREATMENT

Blechnum brackenridgei   comb.

nov.Doodia brackenridgei





Wilkes Exploring

Expedition s.n.





Distribution

Blechnum kermadecense   nom. nov.  

Doodia milnei 

B. milneiLomaria milnei

 



Milne

s.n.



Blechnum kermade-

cense      

   Doodia milnei

Blechnum milnei

                



Distribution.



Perrie & al. • Blechnaceae phylogeny and taxonomy

TAX O N



Version of Record (identical to print version).

Brownlie, G. 

Pacific Sci.  

Brownlie, G. Flore

de la Nouvelle Calédonie et Dépendances       



Brownsey, P.J. & Perrie, L.R. 

Telo p ea

Brownsey, P.J. & Smith-Dodsworth, J.C. New Zealand ferns

and allied plants

Chambers, T.C. Doodia hindii           

Te l opea 

Chambers, T.C.    Stenochlaena

Telope a 

 

Chambers, T.C. & Farrant, P.A.  

Flora of AustraliaFerns, gymnosperms and allied

groups.                



Chambers, T.C. & Farrant, P.A.  Blechnum procerum

capenseNew Zealand

J. Bot.

Blechnum neglectum    

comb. nov. Acrostichum neglectum

Pteridoblech-

num neglectum  





     

 F.M. Bailey s.n. 



Distribution. –

Blechnum reticulatum nom. nov. Lep-

tochilus acuminatus

Pteridoblechnum acuminatum

.

Steenisioblechnum acuminatum  

 B. acuminatum

  







C.T. White 10652



Blechnum 

B. acuminatum

B. acuminatum   

B. occidentale

Distribution. – 

Telmatoblechnumg en. n ov. 

   T. serrulatum

Diagnosis     

      

   









Description     

 

   



 





 



Distribution





Etymolog ytelmat-



Blechnum



Telmatoblechnum indicum

comb. nov. Blechnum indicum   



  



R.G. Coveny 4712   



Distribution



Telmatoblechnum serrulatum 

 comb. nov.  Blechnum serrulatum 

          



 Leblond s.n.  







Distribution

ACKNOWLEDGEMENTS



      

      

    

 









 

 





      

      

   

   



     



LITERATURE CITED



TAX O N

Perrie & al. • Blechnaceae phylogeny and taxonomy

Version of Record (identical to print version).

Chambers, T.C. & Farrant, P.A. Blechnum    

Blumea

Christenhusz, M.J.M., Zhang, X.-C. & Schneider, H. 

   

Phytotaxa

Copeland, E.B.      Genera Filicum  



Cranfill, R. Phylogenetic studies in the Polypodiales (Pterido-

phyta) with an emphasis on the family Blechnaceae



Cranfill, R. & Kato, M. 



     Pteridology in the new

millennium

    

Dawson, M.I., Brownsey, P.J. & Lovis, J.D. 

    New

Zealand J. Bot.



Ebihara, A., Nitta, J.H. & Ito, M. 



PLoS ONE

 

Entwisle, T.J. & Weston, P.H.  

Austral. Syst. Bot. 

    

Gabriel y Galán, J.M., Prada, C., Rolleri, C., Ainouche, A. & Vicent,

M. 

Blechnum

Türk Bot. Derg. = Turk . J. Bot.

Gastony, G.J. & Ungerer, M.C.    



  Amer. J. Bot. 

 

Green, P.S. Flora of Australia

Oceanic islands 1. 

Guindon, S., Dufayard, J.F., Lefort, V., Anisimova, M., Hordijk,

W. & Gascuel, O.    

  

Syst. Biol.

    

Giudice, G.E., Luna, M.L., Carrión, C. & De la Sota, E.R. 

Salpichlaena

   Amer. Fern J.

 

Huelsenbeck, J.P. & Ronquist, F.R. 

Biometrics

Kramer, K.U., Chambers, T.C. & Hennipman, E. 

The families and

genera of vascular plantsPteridophytes and gymnosperms.

 

Larkin, M.A., Blackshields, G., Brown, N.P., Chenna, R.,

McGettigan, P.A., McWilliam, H., Valentin, F., Wallace, I.M.,

Wilm, A., Lopez, R., Thompson, J.D., Gibson, T.J. & Higgins,

D.G.    Bioinformatics



Lehtonen, S. PLoS

ONE 

Löve, A., Löve, D. & Pichi Sermolli, R .E.G.      Cytotaxonomical

atlas of the Pteridophyta

Murray, B.G. & De Lange, P.J. 



New Zealand J. Bot.

     

Nakahira, Y.  A molecular phylogenetic analysis of the family

Blechnaceae, using the chloroplast gene 



Nose, T. & Matsumoto, S.     

Illustrated Flora of ferns

and fern allies of South Pacific Islands.





Palmer, D.D.    Sadleria

Pacific Sci.

Papadopulos, A.S.T., Baker, W. J., Crayn, D., Butlin, R.K., Kynast,

R.G., Hutton, I. & Savolainen, V.  

     Proc. Natl. Acad. Sci. U.S.A. 

   

Parris, B.S. DoodiaFlora of Australia

Ferns, gymnosperms and allied groups.  

.

Passarelli, L., Gabriel y Galán, J.M., Prada, C. & Roller i, C.H.

Blechnum

Grana

    

Posada, D.    Molec.

Biol. Evol.

Rambaut, A. & Drummond, A.J.    



Rolleri, C.H. & Prada, C. 

de Blechnum    

Anales Jard. Bot. Madrid

Rothfels, C. J., Sundue, M.A., Kuo, L.-Y., Larsson, A., Kato, M.,

Schuettpelz, E. & Pryer, K.M.  



Taxo n 

Rothfels, C.J., Larsson, A., Kuo, L.-Y., Korall, P., Chiou, W.-L. &

Pryer, K.M. 

 

Syst. Biol.

Schuettpelz, E. & Pryer, K.M.   

    Ta x o n 

  

Schuettpelz, E . & Pryer, K.M.           

Proc. Natl. Acad. Sci.

U.S. A.

Shepherd, L.D., Perrie, L.R., Parris, B.S. & Brownsey, P.J. 



 trnL-trnF New Zealand

J. Bot.  

Shepherd, L.D., Perrie, L.R. & Brownsey, P.J. 

   

    Asplenium hookerianum. Molec. Ecol. 



Smith, A.R. & Cranfill, R.B. 

Amer. Fern J. 

 



Smith, A.R., Pryer, K.M., Schuet tpel z, E., Korall, P., Schneider,

H. & Wolf, P.G. Taxo n 



Swofford, D.L. 





Tindale, M.D. & Roy, S.K.  

Austral. Syst. Bot.

    

Veillon, J.M. Blechnum francii 

Adansonia



Vences, M., Guayasamin, J.M., Miralles, A. & De La Riva, I.  



 ootaxa

 



Perrie & al. • Blechnaceae phylogeny and taxonomy

TAX O N



Version of Record (identical to print version).

Appendix 1.

 rbcLrps4

rps4 rps4-trnS trnL-trnF

  

 DoodiaPteridoblechnumBlechnum

B. indic umB. serrulatumTelmatoblechnum.

Blechnum. B. amabile B . ambiguum   

Ohlsen B B233  B. appendiculatum   Parr is 12429 B. articulatum

   Perrie & al. BB36  B. australe  

B. austrobrasilianum    B. binervatum

  B. blechnoides  Perrie 3295 & Brownsey 

B. brasiliense            

B. brackenridgei Doodia brackenridgei          Perr ie & al. FJ 2011 123   

 B. camfieldii                Oh lse n BB216    B. cartilagineum 

Bayly 2182  B.  chambersii    Perr ie & Naikatini FJ 2011 260   

 B. chambersii   Wilson 68      

Perrie 4018 & Shepherd  B . chilenseB . colensoi

         Perrie 3572 & Shepherd  B. contiguum             

Perrie NC 2012 31B. corbassonii   Perrie NC 2012 14   

 B. cordatumB. discolorPerr ie

4015 & Shepherd B. dissectum Doodia dissecta Perrie & al.

BB24  B. diversifoliumPerrie NC 2012 157B. doodianum

Doodia heterophylla Ohlsen BB213 B. durum

Perrie 4023 & Shepherd B. eburneum

B. ensiforme B. filiformePerrie 3459 &

Shepherd    B. f luviatile            Wilson 64         

Perrie 4242 & Perrie B. fragile

B. fraseri                    Per rie 3434 & Shephe rd   B. fullagari

B . gibbumPerrie

NC 2012 76   B . glandulosumB. gracile

B . gregsonii  Ohlsen BB228 B. hancockii

B. hirsutumPerrie NC 2012 196B. howe-

anum   B. kermadecense Doodia

milneiParris 12689 B. kunthianum

B. laevigatumB. lenormandii   

Perrie NC 2012 169B. lineare Doodia linearis  Ohlsen

BB14 7  B. loxenseB. magellanicum

B. medium Doodia media 

 Ohlsen BB146     Perrie NC 2012 120

B. melanocaulon       Perrie & Brownsey FJ 2011 182 

B. membranaceum              Per rie 3475 & She pherd  B. microbasis      

 B. microphyllum   B. milnei

     Perrie & Brownse y FJ 2011 165 B. minus    

Wilson 72  Perrie 3323 & al.B. mochaenum

B. molle Doodia mollis      

Parris 12688     B. montanumPerrie 3498

& al.  B. moorei    Perrie NC 2012 146 B. moritzianum

 B. neglectum  Pteridoblechnum neglectum

   Per rie & al. BB62   B. neohollandicum  Doodia

aspera Perr ie & Ohlsen BB113 B. nigrum       

Perrie 3454 & Shepherd  B. norfolkense  Doodia kunthiana

B. norfolkianum    Pa rri s 12358

   B. novae-zelandiae                     Perrie 3129 & Lovis     

B. nudum Wilson 69 B. obtusatum 

Perrie NC 2012 20B. occidentaleB. oceanicum

  Perrie NC 2012 105   B. opacum   

        Perr ie NC 2012 167     B. orientale           Ohlsen & al. BB188 

 Brownsey & Perrie FJ 2011 90      

Perrie NC 2012 204 B. parrisii  Doodia australis   

 Ohlsen BB151    Per rie 3442 & Shepherd   

B. patersoniipatersonii:  Wilson 70  B. patersonii

queenslandicum                   Ohlse n BB144   B. penna-marina        

Wilson 75 Perrie & al. 3497 

B. pilosumPerrie & Brownsey FJ 2011

181     B. polypodioides                  B . procerum          

Wang, F.H., Xing, F.W., Dong, S.Y. & Kato, M. 

Flora

of China  

Wei, R., Schneider, H. & Zhang, X.-C. 

 Diplazium 



Taxon      

Zuloaga, F.O., Morrone, O. & Belgrano, M.J. Catálogo de las

plantas vasculares del Cono Sur (Argentina, sur de Brasil, Chile,

Paraguay y Ur uguay)           



TAX O N

Perrie & al. • Blechnaceae phylogeny and taxonomy

Version of Record (identical to print version).

Appendix 1.



Perrie 3127 & Lovis B. punctulatum  Parr is 12683      

B. reticulatum  Pteridoblechnum acuminatumField

& al. s.n.   B. sampaioanumB. schiedeanum

B. schomburgkii  B. simillimum

 B. spicant

   B. spinulosum    Doodia caudata  Perrie & Ohlsen BB88   

B. stipitellatum         B. subcordatum          

 Perrie NC 2012 29B. tabulare           

B. triangularifolium        Pe rri e 3 416 B. vieillardii  

   Perrie NC 2012 227B. vitta tum Perr ie &

Brownse y FJ 2011 210    B . vulcanicum                  Wilso n 73       

Perrie 3461 & Shepherd   B . wattsiiPerrie &

Ohlsen BB127  B. whelanii    Per rie & al. BB64   B. wurunuran

 Perrie & al. BB34 B. zeelandicum  Doodia squarrosa 

Enright & John s.n. [Perr ie 3517]Brainea. B. insignis

Sadleria. S. cyatheoides

 S. soulayetianaS. squarrosa

Salpichlaena. S. volubilis   

      Stenochlaena. S. milnei         

S. palustri s Ohlsen & Bayly BB197 

 Perrie & Brownsey FJ 2011 84S. tenuifolia   Telmatoblechnum.

T. indicum    Blechnum indicum  Perrie & Ohlsen BB107

 T. serrula tum   Blechnum serrulatum   

Woodwardia. W. areolata

 W. j apon ic a          W. prolifera 

W. radicans 

W. virginica Out-

groups. Matteuccia struthiopterisOnoclea sensibilis

 Onocleopsis hintoniiPentarhizidium

intermedium 

Revision of Cranfillia opaca in New Caledonia

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Apr 2021

Cranfillia opaca is a fern species present in New Caledonia, for which a great variation in morphology is reported. Some other species such as Blechnum deplanchei have been proposed, but those were later synonymized with C. opaca. In this work, we have studied this taxon at both the morphological and the anatomical level, as well as its ecology and geographical distribution. We found differences between C. opaca s.s. and B. deplanchei in the morphology of the sterile and fertile fronds, in several morphoanatomical features such as trichome shape, length, and distribution, and length and/or shape of different structures of sporangia and spores. The distribution of both taxa was also different, but they are sympatric at some localities. The differences identified in this study support the recognition of both taxa, and the combination of Cranfillia deplanchei (Baker) Vázquez Ferreira & Gabriel y Galán, which we propose here. A new taxonomic description is provided for both species.

WHICH IS MORE APPROPRIATE FOR DECIPHERING RECURSIVE PATTERNS IN PLANT RECOVERY ACROSS THE TRIASSIC/JURASSIC AND CRETACEOUS/PALEOGENE BOUNDARIES, NOMOTHETIC OR IDIOGRAPHIC APPROACHES?

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Aug 2022

Keith Berry

Remarkably similar patterns of palynological spikes across the Triassic/Jurassic and Cretaceous/Paleogene boundaries, which are separated by 135 my, beg the question of whether idiographic or nomothetic approaches are better suited to explain these phenomena. Contemporary nomothetic approaches to explaining rapid plant recovery from large-scale natural disasters invoke key traits, such as long-distance dispersal of wind-blown fern spores coupled with whole genome duplication (WGD) (e.g., Baker’s Law), to explain this phenomenon. Detailed investigation of these patterns reveals, however, that there is no universal explanation of this pattern, and that these repetitive patterns must be understood contextually in order to fully grasp nuanced differences between them. Although WGD is frequently linked with the proliferation of ferns or other plant taxa throughout Earth history, for example, it is neither a universal requirement nor always associated with the same key traits (although drought tolerance may be relatively common). Accordingly, idiographic approaches to the study of fern or other plant spikes in geologic history may be more appropriate.

Origin of Hawaiian ferns of the genus Ctenitis (Dryopteridaceae) View supplementary material Origin of Hawaiian ferns of the genus Ctenitis (Dryopteridaceae)

Article

Jul 2022

The Hawaiian Islands are an emblematic field to study evolution, with their very high rates of endemism and spectacular cases of insular speciation. Nevertheless, many Hawaiian lineages still deserve investigation, such as in the fern lineage. In this study, we address the question of the origin of the fern genus Ctenitis, which is present in the archipelago with two endemic species, Ctenitis squamigera and C. latifrons. Using a taxonomic sampling covering the pan-tropical distribution of the genus and three chloroplast DNA regions, we provide evidence that the genus in the Hawaiian Islands originated from a single long-distance dispersal from the Neotropics. This area is less represented than Asia and the South Pacific in the origin of Hawaiian ferns, but a Neotropical origin may be explained by the transportation of spores by tropical storms originating near Central America. Furthermore, the colonization of the Hawaiian Islands is estimated to have occurred between 4 [9-2] and 3 [7-1] mya. This timing is consistent with the ages of all main and extant islands of the archipelago, which already provided habitats for the establishment of the initial Ctenitis colonizer. In turn, this relatively late arrival to the islands and the related potentially low availability of ecological niches may have hampered diversification of the genus beyond the two extant species. ARTICLE HISTORY

Diversity, structure and composition of pteridophyte in varying habitats in Karimun Besar Island, Riau Islands Province, Indonesia

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Full-text available

Oct 2021

Sofiyanti N, Iriani D, Taufik I, Sari M, Irawan A, Syauqi FM. 2021. Diversity, structure and composition of pteridophyte in varying habitats in Karimun Besar Island, Riau Islands Province, Indonesia. Biodiversitas 22: 4847-4856. Biodiversity of small islands, particularly in tropical region, always encourages scientific curiosity. Such curiosity is much greater when the small island of interest has been through various land use changes, such as in Karimun Besar Island. This island is one of thousand islands in Riau Islands Province, Indonesia and poses different land uses. However, studies on the biodiversity of flora in this island have not been recorded, including for pteridophyte or fern members. This study aimed to examine the diversity, floristic structure and composition of pteridophyte at six different study sites in Karimun Besar Island representing varying habitat conditions (i.e., primary forest, secondary forest, rubber plantation, road side, coastal area and mined land. The sampling was collected by using transect method to measure density, dominance and frequency. A total of 32 species of pteridophytes were recorded from the six study sites that belong to two divisions, Polypodiophyta (31 species) and Lycophyta (1 species). The number of species of pteridophyte varied across the six habitats. The highest number of species (21) was found in primary forest, while the lowest (9 species) was found in rubber plantation. The dominant species for each study site were Taenitis blechnoides (Pteridaceae) (rubber plantation), Dicranopteris linearis (Gleicheniaceae) (secondary forest), Sticherus truncates (Gleicheniaceae) (primary forest), Pteridium caudatum (Dennstaedtiaceae) (road side), Achrosticum aureum (Pteridaceae) (coastal area) and Nephrolepis hirsutula (Nephrolepidaceae) (mined land). The result of this study provides the first data of fern in Karimun Besar Island that support pivotal information on the biodiversity of small islands in tropical region.

Deep vicariance and frequent transoceanic dispersal shape the evolutionary history of a globally distributed fern family

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Sep 2022

Premise: Historical biogeography of ferns is typically expected to be dominated by long-distance dispersal, due to their minuscule spores. However, few studies have inferred the historical biogeography of a large and widely distributed group of ferns to test this hypothesis. Our aims are to determine the extent to which long-distance dispersal vs. vicariance have shaped the history of the fern family Blechnaceae, to explore ecological correlates of dispersal and diversification, and to determine whether these patterns differ between the northern and southern hemispheres. Methods: We used sequence data for three chloroplast loci to infer a time-calibrated phylogeny for 154 out of 265 species of Blechnaceae, including representatives of all genera in the family. This tree was used to conduct ancestral range reconstruction and stochastic character mapping, estimate diversification rates, and identify ecological correlates of diversification. Key results: Blechnaceae originated in Eurasia and began diversifying in the late Cretaceous. A lineage comprising most extant diversity diversified principally in the austral Pacific region around the Paleocene-Eocene Thermal Maximum. Land connections that existed near the poles during periods of warm climates likely facilitated migration of several lineages, with subsequent climate-mediated vicariance shaping current distributions. Long-distance dispersal is frequent and asymmetrical, with New Zealand/Pacific Islands, Australia, and tropical America being major source areas. Conclusions: Ancient vicariance and extensive long-distance dispersal have shaped the history of Blechnaceae in both the northern and southern hemispheres. The exceptional diversity in austral regions appears to reflect rapid speciation in these areas; mechanisms underlying this evolutionary success remain uncertain. This article is protected by copyright. All rights reserved.

Revisiting R.H. Tschudy’s fern-spore spike concept 40 years later: Connecting Tschudy’s “disaster taxon” concept with paleopolyploidization in Stenochlaena J. Sm.

Article

Jul 2022
Palynology

Keith Berry

The Cretaceous/Paleogene (K/Pg) boundary fern-spore spike concept was first introduced by R.H. Tschudy 40 years ago and established the precept that ferns are so-called ‘disaster taxa’ that flourish after natural disasters because of the high colonization potential of their wind-blown spores. Among the least understood topics at the crux of contemporary K/Pg boundary fern-spike studies is whether Stenochlaena-like or other stem blechnaceous ferns were among those that flourished after the K/Pg event, contributing to the Laevigatosporites-dominated phase of the fern-spore spike. For instance, dispersed Stenochlaena-like spores – e.g. Polypodiisporites usmensis (van der Hammen) Khan & Martin – first appear in the upper Eocene stratigraphic record of South America, whereas megafossils of Stenochlaena-like ferns are first known from the upper Paleocene strata of this same region. Beyond this traditional focus on taphonomic and taxonomic uncertainty regarding the identity of dispersed spore producers, however, there is further discordance between contemporary paleobotanical and molecular phylogenetic perspectives on the timing of diversification of stem lineages of blechnaceous ferns in relation to the K/Pg boundary. This investigation reconciles these two perspectives by constraining molecular clock divergence time estimates using contemporary fossil data. If this reconciliation is correct, then paleopolyploidization (whole genome duplication or WGD) associated with the origin of the genus Stenochlaena J. Sm. predated evolution of polocytic stomata, a heavily ornamented exospore, and a hemi-epiphytic to epiphytic habit observed in the crown lineage. Critically, this novel perspective elicits a merger of the K/Pg boundary fern spike and K/Pg boundary-WGD concepts,

Molecular phylogenetics and generic taxonomy of Blechnaceae ferns

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