Olaf Werner1, Sergio Pisa1, Susana Rams2, Milagros Saavedra3, Marta Nieto-Lugilde1

& Rosa María Ros1 1Universidad de Murcia, Facultad de Biología, Departamento de Biología Vegetal, Murcia, Spain. E-mail: werner@um.es 2Centro de Magisterio "La Inmaculada", Granada, Spain 3 Instituto de Investigación y Formación Agraria y Pesquera, Consejería de Agricultura y Pesca y Medio Ambiente, Córdoba, Spain

Structure of this talk

Part one: Population genetic structure observed in three common bryophyte species

Bryum argenteum Hedw.

Didymodon vinealis (Brid.) R.H. Zander

Tortula muralis Hedw.

Part two: What do these results tell us about bryophyte biology

Landscape genetics: aims to inform on the interactions between landscape features and evolutionary processes, mainly gene flow and selection.

Within species variation is studied. Important implications for conservation and

evolutionary biology. Frequent species are easier to study:

More samples Higher genetic diversity

Part one: Genetic structure of populations observed in three common bryophyte species 1. Bryum argenteum

Foto: Heike Hofmann

Bryum argenteum distribution

Pisa et al. (2014)

Bryum argenteum

sampling

Distribution of sequence haplotypes in Bryum argenteum

Pisa et al. (2014)

Continental area Asia America Europe Australasia Antarctica Africa sub-Antarctic

Asia - 0.012 n.s. 0.157 n.s. 0.124 (0.0358) 0.383 (0.0003)* 0.060 n.s. 0.100 n.s.

America 0.033 (0.0159)* - 0.135 n.s. 0.049 n.s. 0.479 (0.0001)* 0.027 n.s. 0.062 n.s.

Europe 0.105 (0.0004)* 0.120 (0.0001)* - 0.180 n.s. 0.568 (0.0001)* 0.077 n.s. 0.181 n.s.

Australasia 0.034 n.s.. 0.025 n.s. 0.122 (0.0085)* - 0.565 (0.0002)* 0.114 n.s. 0.016 n.s.

Antarctica 0.186 (0.0001)* 0.210 (0.0001)* 0.272 (0.0001)* 0.251 (0.0001)* - 0.549 (0.0002)* 0.498 (0.0047)*

Africa 0.056 (0.0082)* 0.044 (0.0334)* 0.107 (0.0031)* 0.066 (0.0466) 0.256 (0.0001)* - 0.073 n.s.

Sub-Antarctic 0.096 (0.0007)* 0.088(0.0046)* 0.156 (0.0015)* 0.077 (0.0468) 0.298 (0.0001)* 0.127 (0.0025)* -

n.s. indicates that the test is not significant (P>0.05) * indicates that the test remains significant after the correction of Benjamini and Yekutieli (2001) for multiple tests at the p<0.05 significance level

Valores para FST y NST

Pisa et al. (2014)

Prevalent wind directions in July

Basic conclusions:

Bryum argenteum is capable of frequent (in geological times)long distance dispersal Genetic barriers, even at continental scale, are low

Bryum argenteum Tenerife

Various colonization events

Isolation by distance: p = 0.005

Isolation by altitude: p = 0.0001

Partial Mantel test: p = 0.0001

Bryum argenteum Sierra Nevada and

nearby areas

Isolation by distance: Excluded

Correlation altitude - genetic distance

p < 0,01

Partial Mantel Test, controlling geographic distance

p < 0,01

Pisa et al. (2013)

2. Didymodon vinealis

Distribution of Didymodon vinealis

Didymodon vinealis frecuently covers soil between olive trees. Cabra (Córdoba), 2010.

Didymodon vinealis One of the most frequent mosses in olive plantations

In the presence of mosses there are less weeds, thus reducing the need for herbicides.

The moss cover reduces water evaporation.

In the presence of mosses, harvested olives are less contaminated by mud and as a concequence less soil particles reach the olive press.

Didymodon vinealis ITS tree Didymodon_vinealis_3180_Anelma_5 Didymodon_vinealis_3181_Marmolejo_1 Didymodon_vinealis_3179_Anelma_4 Didymodon_vinealis_3177_Anelma_2 Didymodon_vinealis_3175_Torre_3 Didymodon_vinealis_3174_Torre_2 Didymodon_vinealis_3167_Arroyo_4 Didymodon_vinealis_3165_Arroyo_2 Didymodon_vinealis_3164_Arroyo_1 Didymodon_vinealis_Cabra_2 Didymodon_vinealis_3160_Cabra_1 Didymodon_vinealis_34_16 Didymodon_vinealis_46_9 Didymodon_vinealis_46_11 Didymodon_vinealis_46_19 Didymodon_vinealis_74_5 Didymodon_vinealis_32_1 Didymodon_vinealis_46_5 Didymodon_vinealis_74_11 Didymodon_vinealis_74_14 Didymodon_vinealis_32_17

Didymodon_3168_Arroyo_5 Didymodon_vinealis_3178_Anelma_3 Didymodon_vinealis_3158_Villaharta_1

Didymodon_vinealis_SN_3092 Didymodon_vinealis_34_12

Didymodon_vinealis_SN_3096 Didymodon_vinealis_SN_3097

Didymodon_vinealis_3159_Villaharta_2 Didymodon_vinealis_3169_Castellar_1

Didymodon_vinealis_SN_3106 Didymodon_vinealis_74_7

Didymodon_vinealis_SN_3102 Didymodon_vienalis_3162_Cabra_3

Didymodon_vinealis_SN Didymodon_vinealis_32_8

Didymodon_vinealis_34_11 Didymodon_vinealis_4_10

Didymodon_vinealis_46_16 Didymodon_vinealis_3_7

Didymodon_vinealis_3_18 Didymodon_vinealis_3_11 Didymodon_vinealis_3_13 Didymodon_vinealis_32_9 Didymodon_vinealis_34_8 Didymodon_vinealis_29_4 Didymodon_vinealis_4_13 Didymodon_vinealis_4_8 Didymodon_vinealis_4_16 Didymodon_vinealis_29_1 Didymodon_vinealis_29_10 Didymodon_vinealis_34_2

Didymodon_vinealis_3_15 Didymodon_vinealis_74_1

Didymodon_vinealis_SN_3096 Didymodon_vinealis_SN_3094

Didymodon_vinealis_4_3 Didymodon_vinealis_SN_3094

Didymodon_vinealis_3095_SN Didymodon_vinealis_SN_3093

Didymodon_vinealis_3172_Castallar_4 Didymodon_vinealis_SN_3098

Didymodon_vinealis_SN_3101 Didymodon_vinealis_Ronda

Didymodon_vinealis_SN_3092 Didymodon_vinealis_3170_Castellar_2 Didymodon_vinealis_3176_Anelma_1 Didymodon_vinealis_3166_Arroyo_3 Didymodon_vinealis_MUB13438 Didymodon_vinealis_32_6

Didymodon_vinealis_3163_Cabra_4 Didymodon_vinealis_3102_SN

Didymodon_vinealis_SN_3099 Didymodon_vinealis_SN_3102

Didymodon_vinealis_29_8 Didymodon_vinealis_3171_Castellar_3

Didymodon_vinealis_3182_Marmolejo_2 Didymodon_vinealis_SN_3183

2

31 artificial 25 natural

2 artificial 19 natural

p(Fst) artificial/natural < 0.001

3.Tortula muralis

Grows on rocks, walls and compacted soil

Tortula muralis distribution

Tortula muralis tree cp rps4

Exclusively natural sites

No geographical differentiation: exception Japan

p(FST) natural/artificial < 0.05

Part two: What do these results tell

us on bryophyte biology

Landscape genetics: aims to inform on the interactions between landscape features and evolutionary processes, mainly gene flow and selection.

Are spores and propagules dispersed at long distances?

Byophytes are widely distributed

Generally low differentiation even between distant populations

Because of the reduced size of spores and propagules bryophytes are distributedd like microorganisms

Everything is everywhere,

but the environment selects (Baas Becking, 1934)

Limited gene flow

With high gene flow we need high selection pressure to maintain

distinct populations

The Law of Independent Assortment states that alleles for separate traits are passed independently of one another from parents to offspring

We do not sequence genes with

adaptive advantages How can we use the

corresponding sequence information for landscape genetics?

Genes used for sequencing must be linked to genes with adaptive value

Chromosomes form linkage groups

Chloroplast and mitochondrial DNA don´t recombine

Asexual propagation: axiliary bulbils in Bryum argenteum

Foto George Shepherd

Bryum argenteum is rarely found with sporophytes in many parts of Spain

Didymodon vinealis forms occasionally multicellular gemmae in clusters in leaf axils

The same is true for Didymodon vinealis

Angiosperm life cycle

Selfing in diploid plant

Intragametophytic selfing : antheridia and archegonia on the same plant, monoicous species; Tortula muralis: sperm and ovules are produced by mitosis and therefore identical. Offspring individuals are clones of parent individual

Intergametophytic selfing: mating between separate gametophytes produced by the same sporophyte; Bryum argenteum, Didymodon vinealis. Equivalent to selfing in diploid vascular plants.

Bryophytes:

Several specialist gentotypes?

Genotypes overlapping or not?

frequent generalist genotypes

rare specialist genotypes

frequent generalist genotypes

rare specialist genotypes

Thank you very much for your attention

ACKNOWLEDGEMENTS: to the Spanish Ministry of Science and Innovation (Project CGL2014-52579-R), E.U. FEDER funds and

Fundación SéNeCa