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Octosporopsis nicolai – ein rätselhafter Vertreter aus der Familie der Pyronemataceae

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2014): Octosporopsis nicolai – a mysterious member of the Pyronemataceae. Zeitschrift für Mykologie 80/2: 565-592. Abstract: Several recent finds of Kotlabaea nicolai provided the opportunity to examine this rarely found discomycete in more detail. Molecular data was obtained from various collections. Based on the results of the phylogenetic analysis and the study of morphological features the new genus Octosporopsis U. Lindemann & M. Vega is proposed to accommodate K. nicolai. Zusammenfassung: Auf Basis mehrerer rezenter Funde von Kotlabaea nicolai war es möglich, diesen bisher nur selten nachgewiesenen Discomyceten genauer zu untersuchen. Ein Teil der Aufsammlungen wurde sequenziert, so dass erstmals DNA-Daten für diese Art vorliegen. Aufgrund der Ergebnisse der phylogenetischen Analyse und der Untersuchungen der Mor-phologie wird für Kotlabaea nicolai die neue Gattung Octosporopsis vorgeschlagen.
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© 2014 – Deutsche Gesellschaft für Mykologie
565
Octosporopsis nicolai – ein rätselhafter Vertreter aus der
Familie der Pyronemataceae.


 (2014): Octosporopsis nicolai – a mysterious
member of the Pyronemataceae. 80/2: 565-592.
Key WordsHumaria nicolai, Kotlabaea, Lunularia cruciata -

   Kotlabaea nicolai      
  -
-
  Octosporopsis   
K. nicolai.
Zusammenfassung:Kotlabaea nicolai

     
-
Kotlabaea nicolaiOctosporopsis
Einleitung
-
       -
    Lunularia cruciata 

        -
       
-
        
          -
Humaria nicolai   
       -
         -
-
   


           

: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
566 
Neottiella ricciae 
Neottiella ricciae
           
Kotlabaea nicolai   Kotlabaea

        
-
-
   Kotlabaea    
        

Kotlabaea.


         . 
    K. nicolai mit Kotlabaea
deformis     Kotlabaea   
        
K. nicolai         
        
K. nicolai 
Octosporopsis K. nicolai in Octosporopsis nicolai 

Octosporopsis
 Humaria nicolai

Diagnosis


   
          
       
         
         
           
      
textura intricata/subepidermoidea 
   textura globulosa/angularis    

© 2014 – Deutsche Gesellschaft für Mykologie

          -



-

     
-
-

          -
    -

Etymologie: OctosporopsisOctospora 
        
Octospora   -
 

Octosporopsis nicolai 

Humaria nicolai 
Synonyme: Humarina nicolai 
H. nicolasii 
Kotlabaea nicolai 

nicolasii-
-


: 
     
-
-
-
    
          -

: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 

            
-
-


   :    
          




  -

-

2
-
  
 -
         -

-
     
                 




:     
Pyro-
nemataceae       
O. nicolai
Pyronemataceae
     et
-
           
         -
     -


(
© 2014 – Deutsche Gesellschaft für Mykologie
569
 -
 
-
-
-



    

-

O. nicolai-


-

 


   
O. nicolai




  -
O. nicolai




  - -
O. nicolai




  - -
O. nicolai




  -
O. nicolai




  - -
O. nicolai




   -
O. nicolai




  - -
O. nicolai




  - -
O. nicolai




  - -
: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 



von O. nicolai
O. nicolai-
   Octospora   

       -
O. nicolaiOctospora
(KotlabaeaK. deformis
Scutellinia    
 O. nicolaiKotla-
baeaK. defor-
mis
O. nicolai

 -
-

Fig. 1: 
   O. nicolai 



        Neolecta
Schizosaccharomyces pombe  Marcelleina persoonii 

© 2014 – Deutsche Gesellschaft für Mykologie

© 2014 – Deutsche Gesellschaft für Mykologie
 
Fig. 2:         
OctosporaO. nicolai -
-
    
-
-

© 2014 – Deutsche Gesellschaft für Mykologie

Octosporopsis nicolai 
G: 
 
: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 
Octosporopsis nicolai C:
    D:      E:  
F:J:
  K:        
 
© 2014 – Deutsche Gesellschaft für Mykologie

Octosporopsis nicolai -
H:

  
: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 

O. nicolai basiert auf
   

  -
-




Lunularia cruciata



-
-


-
-




         
         
  



        
         
           
       

      

           


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

  

     
         
  

          
           
         








 meist



          
       
 
          
  

Haare
   
          
      
       

   
          
          

: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 

Octosporopsis nicolai
1a.   

Sagina procumbes 
Barbula fallax Calliergonella cuspidata Lunularia cruciata

Barbula fallaxDicranella staphylina ,
Dicranella varia   Lophocolea bidentata    Lunularia
cruciata        

1c.Lunularia cruciata

1d. 
Lunularia
cruciata         

1e.

    Lunularia cruciata     

1f.

Lunularia cruciata

2.
Aloina aloides
  Barbula unguiculata , Dicranella howei  
 Dicranella  Fissidens bryoides    Pottia starkeana 
  Pseudocrossidium hornschuchianum Riccia
sorocarpa Weissia Lunularia cruciata


Barbula unguiculataDicranella Entosthodon pulchellus
Phascum cuspidatum Pseudocrossidium hornschuchianum
Sphaerocarpos michelii  Lunularia cruciata  
schaftet mit Lamprospora miniata 


4.           
Lunularia cruciataAneura
© 2014 – Deutsche Gesellschaft für Mykologie

pinguis   Aloina aloides Barbula unguiculata Dicranella howei Fissidens
Fossombronia Pottia starckeana, Pseudocrossidium hornschuchianum Phascum
cuspidatum var. piliferum Sphaerocarpus 

-
-
Lunularia cruciata 

6.
Lunu-
laria cruciata         
Lunularia cruciata 

7a.    

Lunularia cruciata 
Lunularia cruciata

   

8.   
-
Lunularia cruciata-



Lunularia cruciata-
        

Kotlabaea deformis
10.


11.
           

12.    


: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 
-

14.-
-


    Octosporopsis nicolai    -
     

-
-


-



 
-






    
           

O. nicolai
-

-
         


   

Kotlabaea nicolai  :      
Kotlabaea“
Humaria nicolai
© 2014 – Deutsche Gesellschaft für Mykologie

          
     Humaria nicolai   
Kotlabaea 

          

       
  Lunularia cruciata     
     
Kotlabaea nicolaiLunularia
cruciata         
O. nicolai nicht an
Lunularia cruciata  
O. nicolai
          
Lunularia cruciata
L. cruciata

 O. nicolai




:-
         O. nicolai
K. deformis 
     -
O. nicolaiK. deformis-
K. deformis


  O. nicolai  K. deformis K. deformis  

   O. nicolai   

   O. nicolai     
       
K. deformisO. nicolai
K. deformis
         

: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 
steht O. nicolaiLunularia
cruciata    
         
    O. nicolai K. deformis
O. nicolai
  Kotlabaea         
 Humaria 
O. nicolaiK. deformis.
  Haare
Sporen-

Sporen-





  

O.
nicolai


(erst
überreif



bei toten




braune



(überreif:

nicht






















braune







mehrfach
septiert
K.
deformis
-





ein-






farbene

(Caroti-

-


farbenen

sehr





nicht
septiert
O. nicolai
   

O. nicolaiOctospora
Octospora (
Octospora NeottiellaLamprospora
       Leucoscypha 
Rhodoscypha  Rhodotarzetta   
O. nicolai
© 2014 – Deutsche Gesellschaft für Mykologie

Rhodoscypha  Rhodotarzetta        
O. nicolai Rhodoscypha
 Rhodotarzetta        
Octospora Neottiella  Lamprospora-

     O. nicolai Leucoscypha leucotricha  
Rhodotarzetta rosea Rhodoscypha ovilla
  O. nicolai       
    
           


Octosporopsis
nicolai
Leucoscypha
leucotricha
Rhodotarzetta
rosea
Rhodoscypha
ovilla






-
-





bis 5 mm im Ø

weiß








-
menium rosafar


weiß





rosa
-
te 

Durchmesser


Ascosporen 

-

-
-







-
meist mit

ein
Nucleus
hyalin





ein Nucleus


mit

-



ein Nucleus
Asci 






-
rorhynch (pers.


: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 
Octosporopsis
nicolai
Leucoscypha
leucotricha
Rhodotarzetta
rosea
Rhodoscypha
ovilla
Paraphysen 

-



-
-




mit












-









Medulla 










Ektales
Exzipulum

-






-







Haare 









-


angeschwollen
-- -


hend

bis mehrfach

-

auslaufend

nicht verbreitert
 bei
Lunularia cruciata



-












auf alten Feuer
stellen-

-
-


-
schen


Nadelhumus
und morschem

fichtenwälder


logie
  

Sommer bis

© 2014 – Deutsche Gesellschaft für Mykologie

Humaria/Kotlabaea
nicolai         Octosporopsis 

        

       


      

       
Octospora/NeottiellaCheilymenia
O. nicolai: -
O. nicolai -

O. nicolai

  

 

          
      
           
 

       
           

  


O. nicolai  
 
von O. nicolai stammt.
O. nicolaiLunularia cruciata-
  

  -
        
von L. cruciata      
         
: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 
L. cruciata – -
-

        
 O. nicolai

O. nicolai
         
            

       
von O. nicolai     
       

besteht O. nicolai


-


 
 < 2
 > 2
 > 2
  > 2
  > 2
 
  
  < 2
  > 2

Schon          O. nicolai offenbar um eine
    



© 2014 – Deutsche Gesellschaft für Mykologie

O. nicolai
            Lunularia
cruciata
           

-

 
von L. cruciataO. nicolai -
-

 L. cruciata

L. cruciata
-
     -

-
 L. cruciata      L. cruciata -
-

 O. nicolaiL. cruciata über-

 
 O. nicolai
     



Phänologie
         

     
(
          


Lunularia cruciata
          
Lunularia cruciata
Lunularia cruciata
: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 

Lunularia cruciata

    -
O. nicolai


        Sept.   
5–22  2
-
O. nicolai bei
          

 O. nicolai    

Lunularia cruciata systematisch nach
O. nicolai


       O. nicolai 
          

         
 
Kotlabaea  
Kotlabaea







     
-
O. nicolai

© 2014 – Deutsche Gesellschaft für Mykologie

O. nicolai


O. nicolai

          
-







Literatur
         

44
-
tus 20
      Leucoscypha leucotricha   
66
Kotlabaea
17
1
           



 SRhodoscypha
Rhodotarzetta16

von Octospora 




 
Rhodoscypha ovilla 

: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
 

8

      
1

 -
67:

 -
6
Lunularia
cruciata-

 Rhodoscypha ovilla Leucoscypha rhodoleuca



40

       Leucoscypha leucotricha  

      -
        

Lunularia vulgaris
. - 
178
 Lunularia cruciata 
Humaria nicolai2
           

-

-
111
 Neottiella ricciae et Bryoscyphus atromarginatus -
22


       
22
© 2014 – Deutsche Gesellschaft für Mykologie
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 ()-

-



28

Kotlabaea nicolai

       -
Octosporopsis
 K. nicolai
combination for K. nicolai is Octosporopsis nicolai 


        
O. nicolaiOctospora.
O. nicolaiKotlabaeaK. deformis 

O. nicolaiK. deformis. In contrast to K. deformis, O. nicolai has
O. nico-
lai

Octosporopsis 


       





Octospora/Neottiella, sometimes of Cheilymenia too.
O. nicolai.

O. nicolai.

 
            
: Octosporopsis nicolai
© 2014 – Deutsche Gesellschaft für Mykologie
592 










LamprosporaOctospora.







            
O. nicolai   L. cruciata 

of L. cruciata     
   O. nicolai
        
L. cruciata.
O. nicolai -
KotlabaeaKotlabaea

... records of known species and new hosts continue to be reported (BeNkerT, 2008;eCk-STeiN & eCkSTeiN, 2009;eCkSTeiN et al., 2014;egerToVÁ et al., 2015). Molecular data have shed light on phylogenetic relationships among the species and have demonstrated narrow host spectra (LiNdeMANN et al., 2014;VegA et al., 2017VegA et al., , 2019egerToVÁ et al., 2018b). Meticulous microscopical observations have also contributed to a better understanding of these parasites by offering new insights about morphological and biological features, such as epispore structures, infected tissues, or gall-forming ability. ...
Article
Döbbeler P, Büschlen A & Ecksein J. 2021. - Octospora bicarpa sp. nov. (Pezizales), the first species of the bryophilous genus Octospora Hedw. with ameroconidia. Ascomycete.org 13(2): 85-91. Octospora bicarpa Döbbeler, Büschlen & Eckstein (Pezizales) is described as a new species of operculate discomycete. It infects the corticolous moss Lewinskya speciosa (Nees) F. Lara, Garilleti & Goffinet (Orthotrichales). The documentation is based on several recent collections from Switzerland. The biotrophic parasite is characterized by small, orange apothecia formed between green leaves in the apical shoot regions. Inconspicuous, almost colourless conidial layers arise near the ascomata in the leaf axils and on the stems. Both morphs originate from the same mycelium, that has infection structures typical for Octospora s. lato, consisting of superficial appressoria and intracellular haustoria. The anamorph also develops independently from the teleomorph. Monoblastic, globose ameroconidia are produced by sterigma-like conidiogenous cells. Octospora bicarpa is the first species of more than 100 recognized octosporaceous fungi that has both ascomata and sporodochia-like conidiomata which produce one-celled conidia.
... Genea Vittad. Lindemann et al. 2014) and Octospora kela-bitiana ( Egertová et al. 2018). Each of the species comprised two genetic lineages that, nevertheless, were relatively weakly diverged and were therefore not treated taxonomically. ...
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Octospora conidiophora is described as a new species, based on collections from South Africa. It is characterised by apothecia with a distinct margin, smooth or finely warted ellipsoid ascospores, stiff, thick-walled hyaline hairs, warted mycelial hyphae and growth on pleurocarpous mosses Trichosteleum perchlorosum and Sematophyllum brachycarpum (Hypnales) on decaying wood in afromontane forests. It is the first species of bryophilous Pezizales in which an anamorph has been observed; it produces long, claviform, curved, hyaline and transversely septate conidia. Three other cryptic species of Octospora were detected using three molecular markers (LSU and SSU nrDNA and EF1α), but these could not be distinguished phenotypically. These are not described formally here and an informal species aggregate O. conidiophora agg. is established for them. The new species and finds of Lamprospora campylopodis growing on Campylopus pyriformis and Neottiella albocincta on Atrichum androgynum represent the first records of bryophilous Pezizales in South Africa. https://mycokeys.pensoft.net/article/34571/
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Eckstein, J., Sochorová, Z. & Janošík, L. 2021. Octospora oscarii spec. nov. (Pezizales), a bryophilous ascomycete on the pleurocarpous moss Pseudotaxiphyllum elegans (Hypnales). – Herzogia 34: 286 –298. The bryophilous ascomycete Octospora oscarii is described as a new species based on collections from Central Germany and the Czech Republic. It grows on the pleurocarpous moss Pseudotaxiphyllum elegans and is characterised by smooth ascospores and an infection on the rhizoids of its host. This is the first account of P. elegans as a host of bryophilous ascomycetes. Phylogenetic analyses of concatenated LSU, SSU and EF1-α gene sequences showed that the studied collections of this species form a well-supported monophyletic lineage differing from all previously studied species of bryophilous Pezizales. Macroscopic and microscopic features of the new species are illustrated by colour photographs and drawings, and are compared to those of the other species of bryophilous Pezizales similarly growing on pleurocarpous mosses.
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Lamprospora angularis sp. nov. is described and illustrated from finds in four different Tenerifan localities. The new species is characterised by a combination of the following features: orange to reddish-orange apothecia with a conspicuous fimbriate to shaggy margin, globose ascospores with distinctive regular areolate ornamentation, and infection of the rhizoids of its bryophyte host Campylopus pilifer. Phylogenetic analysis of SSu, LSu and EF1-α gene sequences show that the studied collections of this species form a well-supported monophyletic clade and clearly differ from all other sequenced species of bryophilous Pezizales. Comparisons have been made with similar members of Lamprospora that infect species of Campylopus, namely L. australis, L. campylopodis and L. verrucispora.
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Lamprospora bulbiformis M.Vega & Janošík, sp. nov., L. gibbosa M.Vega & Janošík, sp. nov. and L. thelespora Martínez-Gil, M.Vega & E.Rubio, sp. nov. are described and illustrated based on live collections from Cyprus, France, Portugal and Spain. Phylogenetic analyses of the concatenated LSU, SSU and EF1-α gene sequences show the studied collections of the three species form well supported monophyletic clades. Lamprospora bulbiformis sp. nov. infects Fissidens viridulus (Sw. ex anon.) Wahlenb., L. gibbosa sp. nov. has F. crassipes Wilson ex Bruch & Schimp. as host and L. the- lespora sp. nov. grows on Cheilothela chloropus (Brid.) Broth. Three Lamprospora De Not. species namely L. tuberculata Seaver, L. tuberculatella Seaver and L. spinulosa Seaver with a slightly similar ascospore ornamentation were described by Seaver from US collections. Results of our studies of their types and additional material collected by Seaver are presented. The host of L. tuberculata is Pleuridium subulatum (Hedw.) Rabenh., that of L. tuberculatella is a species of Weissia Hedw. and that of L. spinulosa is Physcomitrium pyriforme (Hedw.) Bruch & Schimp. It has yet to be proven that species of Lamprospora described from North America also occur in Europe. European collections assigned to any Lamprospora described from North America require revision. It is not unlikely that many or even all of them represent taxa yet to be described. A considerable part of existing literature on bryophilous Pezizales needs to be reevaluated.
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Octospora doebbeleri, a new bryophilous species infecting rhizoids of the acrocarpous moss Dicranoweisia cirrata, is described and illustrated based on collections from the Czech Republic. The delineation of the new species, based on the unique set of morphological characters, is also supported by phylogenetic analysis of LSU and SSU nrDNA and EF1α. Morphologically, O. doebbeleri is characterised by apothecia with a thin margin and ellipsoid, mostly binucleate ascospores ornamented with small isolated warts. Differences from other morphologically similar species of Octospora are discussed.
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The bryophilous ascomycete Octospora svrcekii, belonging to the section Wrightoideae, has so far been reported from only three localities in the world. New collections from Albania, Austria, Croatia, France, Slovakia and Spain have enabled a better understanding of its variability, ecology, distribution and phylogenetic relationships with other taxa within the section Wrightoideae. Octospora svrcekii was always found associated with Cratoneuron filicinum growing in constantly humid habitats (brooks, rivers or waterfalls), on calcareous bedrock. A species description based on both living and dead material is provided and compared with previous observations. A phylogenetic analysis of the section Wrightoideae, performed using the EF1α, SSU rDNA and LSU rDNA loci, revealed that Octospora svrcekii forms a monophyletic group with O. wrightii, O. erzbergeri, O. hygrohypnophila and O. americana, all of which are characterised by subglobose to broadly ellipsoid ascospores ornamented with isolated warts, and infect mosses in the order Hypnales, inducing galls on their rhizoids. Based on the molecular analysis, O. orthotrichi and O. affinis, formerly also considered as members of the section Wrightoideae, do not belong to the group.
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Comparative analysis of molecular sequence data is essential for reconstructing the evolutionary histories of species and inferring the nature and extent of selective forces shaping the evolution of genes and species. Here, we announce the release of Molecular Evolutionary Genetics Analysis version 5 (MEGA5), which is a user-friendly software for mining online databases, building sequence alignments and phylogenetic trees, and using methods of evolutionary bioinformatics in basic biology, biomedicine, and evolution. The newest addition in MEGA5 is a collection of maximum likelihood (ML) analyses for inferring evolutionary trees, selecting best-fit substitution models (nucleotide or amino acid), inferring ancestral states and sequences (along with probabilities), and estimating evolutionary rates site-by-site. In computer simulation analyses, ML tree inference algorithms in MEGA5 compared favorably with other software packages in terms of computational efficiency and the accuracy of the estimates of phylogenetic trees, substitution parameters, and rate variation among sites. The MEGA user interface has now been enhanced to be activity driven to make it easier for the use of both beginners and experienced scientists. This version of MEGA is intended for the Windows platform, and it has been configured for effective use on Mac OS X and Linux desktops. It is available free of charge from http://www.megasoftware.net.
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— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
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Pyronemataceae is the largest and most heterogeneous family of Pezizomycetes. It is morphologically and ecologically highly divers, comprising saprobic, ectomycorrhizal, bryosymbiotic and parasitic species, occurring in a broad range of habitats (on soil, burnt ground, debris, wood, dung and inside living bryophytes, plants and lichens). To assess the monophyly of Pyronemataceae and provide a phylogenetic hypothesis of the group, we compiled a four-gene dataset including one nuclear ribosomal and three protein-coding genes for 132 distinct Pezizomycetes species (4437 nucleotides with all markers available for 80% of the total 142 included taxa). This is the most comprehensive molecular phylogeny of Pyronemataceae, and Pezizomycetes, to date. Three hundred ninety-four new sequences were generated during this project, with the following numbers for each gene: RPB1 (124), RPB2 (99), EF-1α (120) and LSU rDNA (51). The dataset includes 93 unique species from 40 genera of Pyronemataceae, and 34 species from 25 genera representing an additional 12 families of the class. Parsimony, maximum likelihood and Bayesian analyses suggest that Pyronemataceae is paraphyletic due to the nesting of both Ascodesmidaceae and Glaziellaceae within the family. Four lineages with taxa currently classified in the family, the Boubovia, Geopyxis, Pseudombrophila and Pulvinula lineages, form a monophyletic group with Ascodemidaceae and Glaziellaceae. We advocate the exclusion of these four lineages in order to recognize a monophyletic Pyronemataceae. The genus Coprotus (Thelebolales, Leotiomycetes) is shown to belong to Pezizomycetes, forming a strongly supported monophyletic group with Boubovia. Ten strongly supported lineages are identified within Pyronemataceae s. str. Of these, the Pyropyxis and Otidea lineages are identified as successive sister lineages to the rest of Pyronemataceae s. str. The highly reduced (gymnohymenial) Monascella is shown to belong to Pezizomycetes and is for the first time suggested to be closely related to the cleistothecial Warcupia, as a sister group to the primarily apothecial Otidea. None of the lineages of pyronemataceous taxa identified here correspond to previous families or subfamily classifications. Ancestral character state reconstructions (ASR) using a Bayesian approach support that the ancestors of Pezizomycetes and Pyronemataceae were soil inhabiting and saprobic. Ectomycorrhizae have arisen within both lineages A, B and C of Pezizomycetes and are suggested to have evolved independently seven to eight times within Pyronemataceae s.l., whereas an obligate bryosymbiotic lifestyle has arisen only twice. No reversals to a free-living, saprobic lifestyle have happened from symbiotic or parasitic Pyronemataceae. Specializations to various substrates (e.g. burnt ground and dung) are suggested to have occurred several times in mainly saprobic lineages. Although carotenoids in the apothecia are shown to have arisen at least four times in Pezizomycetes, the ancestor of Pyronemataceae s. str., excluding the Pyropyxis and Otidea lineages, most likely produced carotenoids, which were then subsequently lost in some clades (- and possibly gained again). Excipular hairs were found with a high probability to be absent from apothecia in the deepest nodes of Pezizomycetes and in the ancestor of Pyronemataceae s. str. True hairs are restricted to the core group of Pyronemataceae s. str., but are also found in Lasiobolus (Ascodesmidaceae), the Pseudombrophila lineage and the clade of Chorioactidaceae, Sarcoscyphaceae and Sarcosomataceae. The number of gains and losses of true hairs within Pyronemataceae s.str., however, remains uncertain. The ASR of ascospore guttulation under binary coding (present or absent) indicates that this character is fast evolving and prone to shifts.
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Partial sequences of nuLSU rDNA were obtained to investigate the phylogenetic relationships of Pyronemataceae, the largest and least studied family of Pezizales. The dataset includes sequences for 162 species from 51 genera of Pyronemataceae, and 39 species from an additional 13 families of Pezizales. Parsimony, ML, and Bayesian analyses suggest that Pyronemataceae is not monophyletic as it is currently circumscribed. Ascodesmidaceae is nested within Pyronemataceae, and several pyronemataceous taxa are resolved outside the family. Glaziellaceae forms the sister group to Pyronemataceae in ML analyses, but this relationship, as well as those of Pyronemataceae to the other members of the lineage, are not resolved with support. Fourteen clades of pyronemataceous taxa are well supported and/or present in all recovered trees. Several pyronemataceous genera are suggested to be non-monophyletic, including Anthracobia, Cheilymenia, Geopyxis, Humaria, Lasiobolidium, Neottiella, Octospora, Pulvinula, Stephensia, Tricharina, and Trichophaea. Cleistothecial and truffle or truffle-like ascomata forms appear to have evolved independently multiple times within Pyronemataceae. Results of these analyses do not support previous classifications of Pyronemataceae, and suggest that morphological characters traditionally used to segregate the family into subfamilial groups are not phylogenetically informative above the genus level.