1 Benjamin Laenen

Lophocolea heterophylla

Photo B.Laenen

“Large scale phylogenetic patterns of diversification in Bryophytes”

PhD thesis presented byBenjamin Laenen

2 Benjamin Laenen

Thèse déposée en vue de l’obtention du grade de docteur en Sciences biologiques

Université de Liège

“Large scale phylogenetic patterns of diversification in Bryophytes”

Benjamin Laenen

Promoteur : Alain Vanderpoorten

Année 2013

3 Benjamin Laenen

© Copyright. Aux termes de la loi belge du 30 juin 1994, sur le droit d’auteur et les droits voisins, seul l’auteur a le droit de reproduire partiellement ou complètement cet ouvrage de quelque façon et forme que ce soit ou d’en autoriser la reproduction partielle ou complète de quelque manière et sous quelque forme que ce soit. Toute photo-FRSLH�RX�UHSURGXFWLRQ�VRXV�XQH�DXWUH�IRUPH�HVW�GRQF�IDLWH�HQ�YLRODWLRQ�GH�ODGLWH�ORL�HW�GHV�PRGL¿FDWLRQV�XOWpULHXUHV�

“Evolution has no pinnacle and there is no such things as evolutionnary progress”Matt Ridley

“Evolution is within us, around us, between us, and its workings are embedded in the rocks of aeons past”

Richard Dawkins

4 Benjamin Laenen

Summary

In the course of land plant colonization, many once-dominant plant lineages have either gone extinct or been reduced to an ecological role that represents just a shadow of their past glory. The coincidence of WKH�GLYHUVL¿FDWLRQ�RI�HDFK�ODQG�SODQW�OLQHDJH�ZLWK�D�GHFOLQH�LQ�VSHFLHV�QXPEHUV�RI�WKH�SUHYLRXVO\�GRPLQDQW�group has promoted the sequential replacement hypothesis. Bryophytes, which comprise 15-20,000 species of liverworts, mosses, and hornworts, are considered the closest modern relatives of the ancestors to the earliest land plants. The ancient origin of bryophytes and their low level of endemism, together with the close resemblance between fossil and extant taxa raised the hypothesis that bryophytes exhibit a low evolutionary potential. This hypothesis was further circumstantiated by the interpretation of bryophyte species striking range disjunctions in terms of vicariance (ancient vicariance hypothesis, AVH), leading to the consideration of bryophytes as “unmoving, unchanging sphinxes of the past”. In bryophytes, molecular dating analyses evidenced, however, that disjunctions that are spatially consistent with an ancient vicariance hypothesis are much more recent than what that hypothesis would predict, pointing to a major role of dispersal to explain current range disjunctions (Long Distance Dispersal Hypothesis, LDDH). Bryophytes disperse by spores and asexual diaspores, which are assumed to play a complementary role, the former contributing to random LDD and the latter to frequent short-distance dispersal (SDD). Due to fertilization constraints, spores are much more frequently produced by monoicous species than by dioicous ones. Therefore, if, as theory predicts, spores are involved in LDD, range size should also be correlated with sexual condition and monoicous species are expected to exhibit larger ranges than dioicous ones. Inheritability in mating systems and associated life-history traits is, therefore, expected to lead closely related species to exhibit more similar range sizes than distantly related ones, resulting in the intriguing hypothesis that distribution ranges are, similarly, phylogenetically conserved. Alternatively, if range sizes are not explained by the inheritability of mating systems and correlated life-history traits, but result from the asymmetric division of an ancestral range (vicariance), range sizes of sister species are expected to be dissimilar.

In this thesis, I analyzed an 8-genes phylogeny of the liverwort Tree of Life to address the following questions:

%� �Are liverworts, in line with the sequential replacement hypothesis and as the AVH predicts, a slowly evolving lineage? Did they, on the contrary, dynamically take advantage of the new ecological opportunities offered by the emergence of gymnosperm or angiosperm dominated IRUHVWV�DQG�H[SHULHQFH�EXUVWV�LQ�GLYHUVL¿FDWLRQ�UDWHV"�

%� �Can the range of the disjunct distributions exhibited by bryophyte species be explained, as expected by the LDDH, in terms of the heritability of dispersal traits and mating systems, potentially leading to the heritability of range size? Conversely, does asymmetric range inheritance through vicariance, in agreement with the AVH, lead to the absence of phylogenetic signal in range size?

%� �What are the evolutionary trade-offs among mating systems and other life-history traits, shaping the life-history strategies in liverworts?

%� What are the taxonomical consequences of mating system evolution in liverworts?

In full contradiction with views of the group as a ‘sphinx of the past’ with limited evolutionary capacities DQG�VORZ�UDWHV�RI�PROHFXODU�HYROXWLRQ��ZH�VKRZ�WKDW�OLYHUZRUWV�H[SHULHQFHG�VHYHUDO�VKLIWV�LQ�GLYHUVL¿FDWLRQ�rates that peak in the late Cenozoic. Altogether, our results contradict the sequential replacement hypothesis. In line with the interpretation that many adaptations of bryophytes to life in a terrestrial environment, such as water-conducting structures, stomata, and endosymbioses are not the precursors of the ones of vascular plants but instead evolved independently, the results indicate that bryophytes evolved in parallel to vascular plants DQG�DUH�VWLOO�LQ�DQ�DFWLYH�SHULRG�RI�GLYHUVL¿FDWLRQ�

Rejection of the sequential replacement hypothesis points to a much more recent evolutionary origin of extant liverwort diversity that is incompatible with the timing of continental drift and therefore points to a recent long-distance dispersal hypothesis. Despite the strong heritability of dispersal life-history traits, range size was not heritable in the group. In fact, the only trait that correlated to range size was the production of

5 Benjamin Laenen

asexual specialized diaspores, challenging the idea that spores only are involved in LDD and potentially obscuring the correlation between mating systems and range size.

Recurrent transitions from dioicous to monoicous lineages were observed across the liverwort Tree of Life, suggesting that they are adaptive. Although not involved in range size, shifts in mating systems are FRUUHODWHG�WR�D�VXLWH�RI�DGDSWLYH�OLIH�KLVWRU\�WUDLWV�GXULQJ�VSHFL¿F�KDELWDW�WUDQVLWLRQV��

The fast rates of homoplastic evolution in mating systems, as well as the suite of life-history traits that correlates to it, suggest that these characters might be taxonomically misleading and gave an example of application in the genus Leptoscyphus. A more complete re-assessment of liverwort taxonomy in the context of the ‘Liverwort Tree of Life’ project will certainly offer the appropriate framework to propose a revised FODVVL¿FDWLRQ�DQG�DOORZ�D�PRUH�DFFXUDWH�DVVHVVPHQW�RI�FKDUDFWHU�HYROXWLRQ�DQG�ELRJHRJUDSKLFDO�SDWWHUQV�LQ�WKH�group.

6 Benjamin Laenen

Résumé

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$X�FRXUV�GH�FHWWH�WKqVH��M¶DL�XWLOLVp�XQH�SK\ORJpQLH�GHV�KpSDWLTXHV�IDLVDQW�SDUWLH�LQWpJUDQWH�GX�SURMHW�« Liverwort 7UHH�RI�/LIH�ª�SRXU�UpSRQGUH�DX[�TXHVWLRQV�VXLYDQWHV :

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7 Benjamin Laenen

� (Q�WRWDOH�FRQWUDGLFWLRQ�DYHF�O¶LPDJH�GH�©�IRVVLOHV�YLYDQWV�ª�DVVRFLpH�DX[�KpSDWLTXHV��QRXV�PRQWURQV�TX¶HOOHV�RQW�FRQQX�GHX[�SpULRGHV�G¶DFFpOpUDWLRQ�GH�OHXU�GLYHUVL¿FDWLRQ�DYHF�OHV�WDX[�OHV�SOXV�pOHYpV�WURXYpV�DX�&pQR]RwTXH��&HV�UpVXOWDWV�FRQWUHGLVHQW�GRQF�O¶K\SRWKqVH�GX�UHPSODFHPHQW�VpTXHQWLHO�GHV�ÀRUHV�WHUUHVWUHV��(Q�FRQVpTXHQFH��FHUWDLQV�FDUDFWqUHV�GHV�EU\RSK\WHV�WHOV�TXH�GHV�VWUXFWXUHV�YDVFXODLUHV��VWRPDWHV�HW�HQGRV\PELRVHV�QH�VHUDLHQW�SDV�OHV�SUpFXUVHXUV�GH�FHX[�GHV�SODQWHV�YDVFXODLUHV�PDLV�ELHQ�OH�UpVXOWDW�G¶XQH�pYROXWLRQ�HQ�SDUDOOqOH�qui serait toujours en cours.

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� )LQDOHPHQW��OH�WDX[�pOHYp�G¶KRPRSODVLH�GDQV�O¶pYROXWLRQ�GHV�V\VWqPHV�GH�UHSURGXFWLRQ�DLQVL�TXH�OHXU�FRQVpTXHQFH�VXU�OHV�WUDLWV�PRUSKRORJLTXHV�VXJJqUH�TXH�FHV�GHUQLHUV�SRXUUDLHQW�rWUH�WD[RQRPLTXHPHQW�DPELJXV��1RXV�GRQQRQV�LFL�XQ�H[HPSOH�GH�FRQVpTXHQFHV�WD[RQRPLTXHV�G¶XQH�WUDQVLWLRQ�GH�V\VWqPH�GH�UHSURGXFWLRQ�dans le genre Leptoscyphus. Le projet « Liverwort Tree of Life » pose donc les jalons pour construire un cadre VROLGH�HQ�YXH�G¶XQH�UpYLVLRQ�WD[RQRPLTXH�TXL�SHUPHWWUD�XQH�DQDO\VH�SOXV�SUpFLVH�GH�O¶pYROXWLRQ�GHV�FDUDFWqUHV�PRUSKRORJLTXHV�HW�GHV�SDWURQV�ELRJpRJUDSKLTXHV�GDQV�FH�JURXSH�

8 Benjamin Laenen

Acknowledgments

,�ZRXOG�OLNH�WR�WKDQN�¿UVW�P\�VXSHUYLVRU�$ODLQ�9DQGHUSRRUWHQ�IRU�KLV�FRQVWDQW�VXSSRUW�GXULQJ�WKHVH�IRXU�\HDUV�DQG�IRU�WKH�PDQ\�VFLHQWL¿FDOO\�ULFK�GLVFXVVLRQV�ZH�KDG�WRJHWKHU��,�WKDQN�DOVR�-RQ�DQG�%ODQND�6KDZ��LQ�FKDUJH�RI�WKH�/LWRO�SURMHFW��ZKR�OHW�PH�WKH�OHDGHUVKLS�IRU�DQDO\]LQJ�WKH�GLYHUVL¿FDWLRQ�DVSHFWV�LQVLGH�WKH�LPSRUWDQW�Liverwort Tree of Life project. I want also particularly to thank the Pr. Robert Gradstein for his help on the VFRULQJ�RI�FKDUDFWHUV�LQ�OLYHUZRUWV��7KDQNV�DOVR�WR�1LFRODV�'HYRV�ZKR�OHW�PH�SDUWLFLSDWH�LQ�KLV�ZRUN�RQ�WKH�genus Radula��7KDQNV�WR�-HVXV�0XxR]��ZKR�KRVWHG�PH�RQH�PRQWK�LQ�KLV�ODE�LQ�0DGULG��WR�0LFKDHO�6WHFK��IRU�WKH�UHFXUUHQW�VWD\V�,�KDYH�GRQH�LQ�/HLGHQ�8QLYHUVLW\��WR�6WXDUW�0DF'DQLHO��IRU�KRVWLQJ�PH�WKUHH�PRQWKV�LQ�KLV�lab at the University of Florida, and to Laura Forrest, David Bell and David Long for their help during my VWD\�DW�WKH�5R\DO�%RWDQLFDO�*DUGHQ�LQ�(GLQEXUJK��,�DP�JUDWHIXO�WR�WKH�)156�DQG�WKH�)5,$��WR�6<17+(6<6�JUDQWV��16)�DQG�WKH�:%,�

,�DOVR�WKDQN�DOO�WKH�PHPEHUV�RI�WKH�ODE��SHUPDQHQW�RU�WHPSRUDU\��IRU�WKH�VFLHQWL¿F�GLVFXVVLRQV�EXW�DOVR�JRRG�PRPHQWV�ZH�VKDUHG� WRJHWKHU��-DLUR�3DWLxR��9LUJLQLH�+XWVHPpNHUV��6LOYLD�&DOYR��0DQXHO�*LO�/RSH]��5XEHQ�0DWHR��0HUFLD�6LOYLD��/DXUHQW�*RK\�DQG�)ORULDQ�=DQDWWD�

7KDQNV�WR�P\�IDPLO\�IRU�WKHLU�VXSSRUW��HVSHFLDOO\�WR�P\�JRGPRWKHU�-DQLFN��ZKR�GHVSLWH�WKH�GLI¿FXOW\�RI�OLIH�KDV�DOZD\V�EHHQ�VXSSRUWLYH��7KDQNV�DOVR�WR�P\�IDWKHU�DQG�PRWKHU�LQ�ODZ��/XF�DQG�<YHWWH�IRU�WKHLU�JHQHUDO�LQWHUHVW�and the constant care they provided.

)LQDOO\��,�ZDQW�WR�WKDQN�$XUpOLH�'pVDPRUp��ZLWK�ZKRP�,�KDYH�VKDUHG�DOO�WKH�JRRG�DQG�EDG�PRPHQWV�GXULQJ�those fours years, and with whom I want to share the future ones.

Benjamin

9 Benjamin Laenen

Table of contents

• Summary (English – Français)

• Acknowledgments

• General introduction

• Chapter one: “Old but still active: bryophytes are not a sphinx of the past”

• Chapter two: “Range size heritability in liverworts: does sex really matter?”

• Chapter three: “Evolution of sexual systems, dispersal strategies and habitat selection in the liverwort genus Radula”

• Chapter four: “Striking autapomorphic evolution in Physotheca J.J.Engel & Gradst. (Marchantiophyta: Lophocoleaceae) blurred its actual relationships with Leptoscyphus Mitt.”

• Discussion

• List of publications

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