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Manuel Pimentel Pereira, PhD




    Taxonomy and Phylogeography of
 complex species groups. The case of the
      European sweet vernal grasses
  (Anthoxanthum L., Poaceae, Pooideae)



                                           1
Manuel Pimentel Pereira, PhD
  Research experience:




                         - Numerical Taxonomy
                         (Multivariate analysis)




                         - Molecular Taxonomy
                                (RAPDs)




                                                   2
Manuel Pimentel Pereira, PhD
    Research experience:




-Ecological study of Galician peatlands

-Collaboration in fieldwork
                                          -Ecological study of some Galician
-Vegetation inventory                     populations of Coenagrion
                                          mercuriale. Valoration of anthropic
-Evaluation of the observed anthropic     impacts
impacts
                                                                          3
Manuel Pimentel Pereira, PhD




    Taxonomy and Phylogeography of
 complex species groups. The case of the
      European sweet vernal grasses
  (Anthoxanthum L., Poaceae, Pooideae)



                                           4
Genus Anthoxanthum L in Europe.
                        Introduction


                            • Inflorescence a spiciform panicle



                            • Many laterally-compressed spikelets


                            • Glumes unequal, membranous, carinate and
                            mucronate

                            • Three flowers. The upper one fertile, the
                            lower ones reduced to aristate lemmae


                            • Bi-staminate fertile flower. Without lodicles

Drawing: D. Romero


                                                                              5
Genus Anthoxanthum L in Europe.
                                      Introduction
80N



60N



40N



20N



 0



20S



40S



60S



80S
      160W 140W 120W 100W 80W   60W   40W   20W     0    20E   40E    60E    80E   100E 120E   140E 160E
                          Native distribution of genus Anthoxanthum L.s. str .
                          Invaded range of Genus Anthoxanthum L. s . str .


                                                                                                           6
Genus Anthoxanthum L in Europe.
                            Introduction
• Seven Anthoxanthum species are part of the European and
Macaronesian Flora:

      • Anthoxanthum amarum Brot.

      • Anthoxanthum maderense Teppner

      •Anthoxanthum odoratum L. s. lat. complex

             Anthoxanthum odoratum L.
             Anthoxanthum alpinum Löve & Löve

      • Anthoxanthum gracile Bivona

      •Anthoxanthum aristatum/Anthoxanthum ovatum complex
             Anthoxanthum aristatum Boiss.
             Anthoxanthum ovatum Lag.


                                                            7
Genus Anthoxanthum L in Europe.
                           Introduction
Genus Anthoxanthum presents a high Taxonomic and Phylogeographic
interest:
                                                A. odoratum
      1.- Unclear phylogenetic and taxonomic relationships with close genera
             A. alpinum

      2.- The taxonomical status of some of the European species has been
      widely discussed and is still to be established. The great variety of
      ploidy levels that can be found throughout the genus adds complexity
      to the matter




      A. alpinum                               A. odoratum



                                                                               8
Genus Anthoxanthum L in Europe.
                          Introduction
Genus Anthoxanthum presents a high Taxonomic and Phylogeographic
interest:




           A. aristatum                  A. ovatum


                                                                   9
Genus Anthoxanthum L in Europe.
                               Introduction
Genus Anthoxanthum presents a high Taxonomic and Phylogeographic
interest:
        1.-   Unclear phylogenetic and taxonomic relationships with close genera
 A. gracile
                   A. amarum

       2.- The taxonomical status of some of the European species has been
       widely discussed and is still to be established. The great variety of
       ploidy levels that can be found throughout the genus adds complexity
       to the matter

        3.- Given the distribution of some of these species, genus Anthoxanthum
        presents a high phylogeographical interest



                                         A. maderense




                                                                                   10
Genus Anthoxanthum L in Europe.
                        Introduction
Genus Anthoxanthum presents a high Taxonomic and Phylogeographic
interest:
The A. aristatum/A. ovatum complex




                                                                   11
Genus Anthoxanthum L in Europe.
                         Introduction
Genus Anthoxanthum presents a high Taxonomic and Phylogeographic
interest:
  The A. odoratum s. lat. complex




                                                                   12
Genus Anthoxanthum L in Europe.
                                 Research aims

1. To clarify, through morphological, anatomical and molecular methods, the
   taxonomy of the European sweet vernal grasses. A special effort was made on
   the A. odoratum/A. alpinum and the A. aristatum/A. ovatum complexes
2. To calibrate the usefulness and reliability of the traditionally used traits in the
   differentiation of the European species of Genus Anthoxanthum L. Also, to
   analyse the possible taxonomic consequences of ploidy differences among taxa
3. To analyse the influence of the environmental factors on the morphological
   variation observed on some of the studied populations
4. To elaborate a taxonomic proposal for the European species of Anthoxanthum.
5. To unravel the evolutionary differentiation and the phylogeographic patterns of
   some populations of the European sweet vernal grasses




                                                                                   13
The European sweet vernal
grasses. A morphometric study




                                14
Genus Anthoxanthum L in Europe.
             Morphometric study. Methodology
• Can we unambiguously characterize the Anthoxanthum species using
macromorphology and leaf anatomy?




                        •1787 plants from 60 European populations were
                        used

                            • Macromorphology: 1567 plants

                            • Leaf anatomy: 220 plants


                        • A. nivale K. Schum. and A. aethiopicum I.
                        Hedberg

                                                                      15
Genus Anthoxanthum L in Europe.
                         Morphometric study. Methodology


                                                                   29 anatomical characters
   30 macromorphological characters



13 quantitative traits      17 qualitative traits


                                                    7 quantitative traits      22 qualitative traits




                          Gower coefficient for mixed data

                               Cluster analysis
                               Principal coordinates analysis

                          Information radius. Cluster analysis

                                                                                                   16
Genus Anthoxanthum L in Europe.
                         Morphometric study. Methodology


   30 macromorphological characters



13 quantitative traits      17 qualitative traits


                                                    Contingency tables. Lambda coefficient

                                                    Principal coordinates analysis

                                                    Cluster analysis


     Exploratory analysis
     Principal components analysis

     Classification discriminant analysis
     Multivariate analysis of variance

                                                                                             17
Genus Anthoxanthum L in Europe.
                             Morphometric study. Results
         1. Inflorescence lenght
         2. Leaf length
         3. Leaf width
                                                      • Only the Mediterranean diploid A. gracile
                                                      and the high Iberian polyploid A. amarum
                                                      can be easily distinguished based on
                                                      morphological characters


                                                      A. amarum is partially characterized by
                                                      vegetative traits

                                                      A. gracile is clearly characterized by floral
                                                     traits

                                     1. Awn length
                                     2. Sterile floret length
                                     3. Sterile floret width

• Principal components analysis performed with macromorphological quantitative data. The two first
axis accounted for 35.4% of the variance
                                                                                               18
Genus Anthoxanthum L in Europe.
                            Morphometric study. Results

                                                          The differentiation between annuals and
                                                         perennials is much clearer when we
                                                         consider the whole macromorphological
                                                         dataset

                                                         The MANOVA analysis indicated that
                                                         inflorescence and leaf length were
                                                         important to differentiate between
                                                         annuals and perennials


                                                          Morphological variation in A. odoratum
                                                         is, to some extent, geographically
                                                         structured. Pubescence characters are the
                                                         basis of such structure


•Principal coordinates analysis performed with macro-morphological data (Gower coefficient)
•Cluster analysis performed with qualitative macro-morphological data. IP, Iberian
Peninsula; SS, South of Scandinavia
                                                                                              19
Genus Anthoxanthum L in Europe.
                             Morphometric study. Results


     A. odoratum                                       A. maderense                others (7.4%)
                   A. alpinum (4.1%)
                                       others (3.9%)




                                                                               A. odoratum (26.6%)
                                                        A. maderense (66%)
              A. odoratum (92%)
•Classification discriminant analysis performed with macromorphological quantitative data



    A clear differentiation between A. odoratum and A. alpinum was detected

   A. maderense characterization is not possible based on the selected characters

                                                                                               20

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PresentacióN Renovada Pimentel.20

  • 1. Manuel Pimentel Pereira, PhD Taxonomy and Phylogeography of complex species groups. The case of the European sweet vernal grasses (Anthoxanthum L., Poaceae, Pooideae) 1
  • 2. Manuel Pimentel Pereira, PhD Research experience: - Numerical Taxonomy (Multivariate analysis) - Molecular Taxonomy (RAPDs) 2
  • 3. Manuel Pimentel Pereira, PhD Research experience: -Ecological study of Galician peatlands -Collaboration in fieldwork -Ecological study of some Galician -Vegetation inventory populations of Coenagrion mercuriale. Valoration of anthropic -Evaluation of the observed anthropic impacts impacts 3
  • 4. Manuel Pimentel Pereira, PhD Taxonomy and Phylogeography of complex species groups. The case of the European sweet vernal grasses (Anthoxanthum L., Poaceae, Pooideae) 4
  • 5. Genus Anthoxanthum L in Europe. Introduction • Inflorescence a spiciform panicle • Many laterally-compressed spikelets • Glumes unequal, membranous, carinate and mucronate • Three flowers. The upper one fertile, the lower ones reduced to aristate lemmae • Bi-staminate fertile flower. Without lodicles Drawing: D. Romero 5
  • 6. Genus Anthoxanthum L in Europe. Introduction 80N 60N 40N 20N 0 20S 40S 60S 80S 160W 140W 120W 100W 80W 60W 40W 20W 0 20E 40E 60E 80E 100E 120E 140E 160E Native distribution of genus Anthoxanthum L.s. str . Invaded range of Genus Anthoxanthum L. s . str . 6
  • 7. Genus Anthoxanthum L in Europe. Introduction • Seven Anthoxanthum species are part of the European and Macaronesian Flora: • Anthoxanthum amarum Brot. • Anthoxanthum maderense Teppner •Anthoxanthum odoratum L. s. lat. complex Anthoxanthum odoratum L. Anthoxanthum alpinum Löve & Löve • Anthoxanthum gracile Bivona •Anthoxanthum aristatum/Anthoxanthum ovatum complex Anthoxanthum aristatum Boiss. Anthoxanthum ovatum Lag. 7
  • 8. Genus Anthoxanthum L in Europe. Introduction Genus Anthoxanthum presents a high Taxonomic and Phylogeographic interest: A. odoratum 1.- Unclear phylogenetic and taxonomic relationships with close genera A. alpinum 2.- The taxonomical status of some of the European species has been widely discussed and is still to be established. The great variety of ploidy levels that can be found throughout the genus adds complexity to the matter A. alpinum A. odoratum 8
  • 9. Genus Anthoxanthum L in Europe. Introduction Genus Anthoxanthum presents a high Taxonomic and Phylogeographic interest: A. aristatum A. ovatum 9
  • 10. Genus Anthoxanthum L in Europe. Introduction Genus Anthoxanthum presents a high Taxonomic and Phylogeographic interest: 1.- Unclear phylogenetic and taxonomic relationships with close genera A. gracile A. amarum 2.- The taxonomical status of some of the European species has been widely discussed and is still to be established. The great variety of ploidy levels that can be found throughout the genus adds complexity to the matter 3.- Given the distribution of some of these species, genus Anthoxanthum presents a high phylogeographical interest A. maderense 10
  • 11. Genus Anthoxanthum L in Europe. Introduction Genus Anthoxanthum presents a high Taxonomic and Phylogeographic interest: The A. aristatum/A. ovatum complex 11
  • 12. Genus Anthoxanthum L in Europe. Introduction Genus Anthoxanthum presents a high Taxonomic and Phylogeographic interest: The A. odoratum s. lat. complex 12
  • 13. Genus Anthoxanthum L in Europe. Research aims 1. To clarify, through morphological, anatomical and molecular methods, the taxonomy of the European sweet vernal grasses. A special effort was made on the A. odoratum/A. alpinum and the A. aristatum/A. ovatum complexes 2. To calibrate the usefulness and reliability of the traditionally used traits in the differentiation of the European species of Genus Anthoxanthum L. Also, to analyse the possible taxonomic consequences of ploidy differences among taxa 3. To analyse the influence of the environmental factors on the morphological variation observed on some of the studied populations 4. To elaborate a taxonomic proposal for the European species of Anthoxanthum. 5. To unravel the evolutionary differentiation and the phylogeographic patterns of some populations of the European sweet vernal grasses 13
  • 14. The European sweet vernal grasses. A morphometric study 14
  • 15. Genus Anthoxanthum L in Europe. Morphometric study. Methodology • Can we unambiguously characterize the Anthoxanthum species using macromorphology and leaf anatomy? •1787 plants from 60 European populations were used • Macromorphology: 1567 plants • Leaf anatomy: 220 plants • A. nivale K. Schum. and A. aethiopicum I. Hedberg 15
  • 16. Genus Anthoxanthum L in Europe. Morphometric study. Methodology 29 anatomical characters 30 macromorphological characters 13 quantitative traits 17 qualitative traits 7 quantitative traits 22 qualitative traits Gower coefficient for mixed data Cluster analysis Principal coordinates analysis Information radius. Cluster analysis 16
  • 17. Genus Anthoxanthum L in Europe. Morphometric study. Methodology 30 macromorphological characters 13 quantitative traits 17 qualitative traits Contingency tables. Lambda coefficient Principal coordinates analysis Cluster analysis Exploratory analysis Principal components analysis Classification discriminant analysis Multivariate analysis of variance 17
  • 18. Genus Anthoxanthum L in Europe. Morphometric study. Results 1. Inflorescence lenght 2. Leaf length 3. Leaf width • Only the Mediterranean diploid A. gracile and the high Iberian polyploid A. amarum can be easily distinguished based on morphological characters A. amarum is partially characterized by vegetative traits A. gracile is clearly characterized by floral traits 1. Awn length 2. Sterile floret length 3. Sterile floret width • Principal components analysis performed with macromorphological quantitative data. The two first axis accounted for 35.4% of the variance 18
  • 19. Genus Anthoxanthum L in Europe. Morphometric study. Results The differentiation between annuals and perennials is much clearer when we consider the whole macromorphological dataset The MANOVA analysis indicated that inflorescence and leaf length were important to differentiate between annuals and perennials Morphological variation in A. odoratum is, to some extent, geographically structured. Pubescence characters are the basis of such structure •Principal coordinates analysis performed with macro-morphological data (Gower coefficient) •Cluster analysis performed with qualitative macro-morphological data. IP, Iberian Peninsula; SS, South of Scandinavia 19
  • 20. Genus Anthoxanthum L in Europe. Morphometric study. Results A. odoratum A. maderense others (7.4%) A. alpinum (4.1%) others (3.9%) A. odoratum (26.6%) A. maderense (66%) A. odoratum (92%) •Classification discriminant analysis performed with macromorphological quantitative data A clear differentiation between A. odoratum and A. alpinum was detected A. maderense characterization is not possible based on the selected characters 20