NSF Tree of Life Program | TOLKIN

LiToL: Assembling the Liverwort Tree of Life

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Leafy II

Bazzania novae-zelandiae (photo by J. Braggins)

This clade requires more intense focus in order to understand morphological synapomorphies and character evolution within it.  Currently, we characterize this group primarily by the lack of characteristics found in Leafy I.  The clade in general can be differentiated from Leafy I by the possession of ventral branching, also, any complicate-bilobed taxa have a larger ventral lobe.

The Leafy II clade can be further broken into sub-clades.  Figure 9 (Forrest et al., 2006) illustrates these:  clades A, B, and C, Schistochila, and Ptilidium plus Neotrichocolea.

 

Figure 9. Leafy clade II (based on the topology from the 5 partition Bayesian analysis of 5 loci including chloroplast, nuclear, and mitochondrial DNA). Numbers above the branches are Maximum Parsimony bootstrap values/homogeneous Bayesian posterior probabilities (PPs)/5-partition Bayesian PPs/14-partition Bayesian PPs.

Clade A, containing the Lepidoziaceae, Lepicoleaceae (in part), Herbertaceae, and Geocalycaceae (in part; see Classification) has no currently recognized defining feature. However, the Lepidoziaceae, Lepicoleaceae, and Herbertaceae (Acromastigum through Triandrophyllum, Fig. 9) are united by incubous leaf insertion, 2, 3 or 4 parted leaves, and endogenous branches.  Plagiochila (Plagiochilaceae) and the Geocalycaceae (Lophocolea and Chiloscyphus, Fig. 9) have succubous leaf insertion. 

Lepicolea scolopendra (photo by J. Braggins)

Trichocolea molissima (photo by J. Braggins)

Clade B, containing the Scapaniaceae, Jungermanniaceae (in part), Cephaloziellaceae, Cephaloziaceae, and Adelanthaceae (see Classification), has no currently recognized synapomorphy, however, all members have succubous or transverse leaf insertion.

Scapania nemorea with sporophyte (photo by L. Zhang)

Gyrothyra underwoodiana with sporophyte (photo by L. Zhang)

 

Clade C, containing the Gymnomitriaceae (in part), the Balantiopsidaceae, the Acrobolbaceae, some members of the Jungermanniaceae, and several other liverworts from an assemblage of families (see Classification), is defined by the presence of a marsupium or some type of geotropic growth.  All members except Calypogeia (Calypogeiaceae) have succubous leaf insertion.

The Ptilidium/Neotrichocolea clade is only weakly supported as a member of Leafy II in some analyses; in others, it falls to an unresolved position, grouping neither with Leafy I or Leafy II. Neotrichocolea has ventral lobules that are smaller than its dorsal lobes, however, Ptilidium does not possess ventral lobules, and both taxa lack other characteristics associated with Leafy I

Schistochila appendiculata (photo by C. Davis)

Haplomitriaceae and Treubiaceae
Complex Thalloids and the Blasiaceae
Simple thalloids I
Simple thalloids II plus the Pleuroziaceae
Leafy I
Leafy II

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