Cryptogamie, Bryologie, 2010, 31 (2): 125-133 © 2010 Adac. Tous droits réservés

The systematic position of Pedinophyllum (: )

Kathrin FELDBERGa , Ji ÷ íVÁ~ A b , Rui-Liang ZHUc & Jochen HEINRICHSa*

a Georg August University Göttingen, Department of Systematic Botany, Albrecht von Haller Institute of Sciences, Untere Karspüle 2, 37073 Göttingen, Germany.

b Charles University, Department of Botany, Benátská 2, 128 01 Praha 2, Czech Republic.

c Department of Biology, East China Normal University, 3663 Zhong Shan North Road, Shanghai 200062, China.

(Received 12 December 2009, accepted 5 February 2010)

Abstract – Phylogenetic analyses of an rbcL sequence dataset indicates that the generitype Pedinophyllum interruptum nests within the Plagiochilaceae. Previous reports of Pedinophyllum truncatum in the Adelanthaceae or Jamesoniellaceae are likely based on a misidentification of nipponica.

Adelanthaceae / Jamesoniellaceae / Pedinophyllum / Plagiochilaceae / rbcL

INTRODUCTION

The Plagiochilaceae are made up of the species-rich genus (Dumort.) Dumort. and several satellite genera such as Chiastocaulon Carl, Dinckleria Trevis., Pedinophyllopsis R.M. Schust. et Inoue, Pedinophyllum (Lindb.) Lindb. and Plagiochilion S. Hatt. (Crandall-Stotler et al., 2009). However, the circumscription and systematic position of Pedinophyllum (Lindb.) Lindb. is still subject to controversy. This genus shares the basally-inflated, laterally-compressed, bilabiate perianths with other representatives of Plagiochilaceae but differs in its autoicous condition. Based on nrITS and cp DNA rps 4 sequences of a single specimen, Groth & Heinrichs (2003), Heinrichs et al. (2006) and Hentschel et al. (2007) confirmed the placement of the generitype Pedinophyllum interruptum (Nees) Kaal. in Plagiochilaceae. A second species, Pedinophyllum truncatum (Steph.) Inoue, was identified as an element within the Jamesoniellaceae (= Adelanthaceae, Feldberg et al., 2010) by He-Nygrén et al. (2004, 2006) based on a cp DNA rbcL, trn L-F and rps 4 sequence set. Heinrichs et al. (2005) and Hentschel et al. (2006) confirmed the position of Pedinophyllum truncatum in Adelanthaceae using the same sequences. The presence of the two species in different suborders is surprising considering the alternative treatment of

* Correspondence and reprints: [email protected] 126 K. Feldberg, J. Vá≈ a, R.-L. Zhu & J. Heinrichs the taxa as subspecies of a single species (Inoue, 1958). Crandall-Stotler et al. (2009) assumed a misidentification of the Pedinophyllum truncatum DNA voucher; however, this hypothesis has not yet been verified using further accessions. Here we produce rbcL sequences of further Pedinophyllum inter- ruptum specimens, and provide evidence that Pedinophyllum truncatum is sometimes confused with members of the Syzygiella autumnalis (DC.) Feldberg et al./ S. nipponica (S.Hatt.) Feldberg et al. complex.

MATERIALS AND METHODS

DNA extraction, PCR amplification and sequencing were carried out as described in Hentschel et al. (2006). In case no product was detected a second (nested) PCR was performed using the forward primer rbcL-210-F and the reverse primer rbcL-1200-R (Gradstein et al., 2006). Several new rbcL sequences of Pedinophyllum interruptum and putative Pedinophyllum truncatum were included in an alignment of more than 500 published rbcL sequences of liverworts including those from the Adelanthaceae dataset of Feldberg et al. (2010). Maximum parsimony analyses allowed for an identification of closely related taxa. Based on these analyses (data not shown) a subset of 62 sequences was compiled for further analyses (Table 1). The alignment was constructed using Bioedit (Hall, 1999). Lacking parts of sequences were coded as “N” (A, C, G or T). Two representatives of Schistochilaceae were designated as outgroups based on the phylogenies of He-Nygrén et al. (2006) and Heinrichs et al. (2007). Further representatives of Jungermanniales in the circumscription of Heinrichs et al. (2005) formed the ingroup. Maximum likelihood (ML) and parsimony (MP) analyses of the dataset were carried out using PAUP* (Swofford, 2003). The MP analysis was performed with the following options implemented: heuristic search mode with 100 random-addition sequence replicates, tree bisection-reconnection branch swapping (TBR), MULTrees option on, and collapse zero-length branches off. All characters were treated as equally weighted and unordered. Bootstrap support (BS) values were estimated by calculating 500 bootstrap replicates (Felsenstein, 1985). To decide on the nucleotide substitution model with the smallest number of parameters that best fits the data the program Modeltest (Posada and Crandall, 1998) was used. The Akaike information criterion (Akaike, 1974) selected the GTR model (Tavaré, 1986) with gamma shape parameter foramong site variation and proportion of invariable sites. This model was implemented in ML analyses carried out as heuristic searches. Branching confidence was assessed using 100 bootstrap resamplings in ML analysis using a neighbor joining tree as starting tree and allowing for 2000 rearrangements under the GTR + I + G model. Clade support was also assessed by bootstrapping with 200 replicates, accomplished with GARLI version 0.96 (Zwickl, 2008) using the default search parameters. Bootstrap percentage values above 70 were regarded as good support (Hillis & Bull, 1993).

RESULTS

Three hundred and twenty one of the 1122 aligned characters of the rbcL gene were parsimony-informative, 105 autapomorphic and 696 constant. The strict The systematic position of Pedinophyllum (Marchantiophyta: Jungermanniales)127 Acc .- N o. Y 4 622 85 Y 6 9999 0 Y 4 622 9 2 Y 4 623 14 Q 3 1 2 4 77 Q 3 1 2 49 Q 3 1 2 49 Q 3 1 2 5 02 Q 3 1 2 5 0 EF 5 0367 4 AY 4 622 91 AY 149851 AY 4 622 94 AY 5 073 9 7 AY 4 62307 AY 5 07 4 0 9 DQ 3 1 2 4 76 DQ 026 58 2 DQ 194 07 4 G en Ba nk s eq u en c e K ingdom Z e a l nd Z e a l nd Z e a l nd Z e a l nd a ll SA fo r N e w G e r m a n y EF 5 0367 P e ru A ustr a li aA in c l u ded a r e L] ( GOET ) N e w r bc ( GOET ) [ ( GOET ) G e r m a n y DQ 3 1 2 5 0 8 ( ABSH ) USA 1 3777 0 1 7 84 1 07 ( H ) F inl a nd 0 1 6 18 n u m b e rs ( GOET ) U ni t ed B ry o V o u c he r O igin ( GOET ) G e r m a n y DQ 3 1 2 481 3 51 ]U 14 6 9 ( GOET ) B u lg a r i aD ( GOET ) N o rw a y DQ 3 1 2 491 ( GOET ) S lo v a ki aD a n l y e s . B ry o V e rw imp ( GOET ) A ustr i aD ( GOET ) Pa n a m aD ( EGR ) Ma d a g s ca r GQ 9 002 95 CANB 63 9918) & no. ( GOET ) I ndone s i aD Z ML 07 5 3 0 1159 0 458 0 8 6 4 ( H ) A r gen t in aA ( JE ) G e r m a n y DQ 3 1 2 49 0 ( GOET ) A ustr i aA ( GOET ) B elgi u m AY 6 99994 4 3 9 0 15 7 8 P iippo ( GOET ) Ma d a g s ca r FJ 98494 0 ( H ) A ustr a li aA G en ba nk 2 11 6 - 41 0 ( H ) N e w H ein r i c h s W il s on [ r ef. f r om ( GOET ) N e w a l. s .n. ( H ) F inl a nd & a l. a nd s .n. B ry o B ry o B ry o & B ry o & 1 02 4 0 4 3 88 3 8 26 e t ( HIRO ) Ja p a n AY 6 99991 C r a nd ll- S t o le 5 637 a a l. e t R 9 6 s .n. NH 48 0 4 94 6 1 / A, 8918 85 20 & (lo a n e t 5 6 55 84 MP 20 85 3 1 7 54 2 N ep a l AY 4 62300 in P eng e- N y g r én s eq u en c e . H en ts c hel H en ts c hel C ur no w V oij t ko R o t he r G r a d st ein A h H en ts c hel Da u phin us ed mole c u l a r fo r Ha mlin Sc h ust . G lenn y Sc h ust . G lenn y M i tt . E ngel P e a rs on) s pe c imen Sc hiffn. R . M e t R . M ) T r e v i s . Sc h ä fe - V rw imp P e a rs on e t a l. M i tt . H yv önen G r olle Ta y lo r ) D u mo rt . W heele r e t ) G r olle e t G r olle Ca r l K ur i t a a l. L oe s ke ( H ook.) f. L ind b . H ein r i c h s Ta y lo r ) Ta x on E ngel V o u c he r N ee s F r a hm N ee s H ein r i c h T r e v i s . L ong N ee s M ein u nge r e t Sc hiffn. C o r d aH D u mo rt . H ein r i c h s e t ( Ca rr ing t on 1 : S p ru c e F r a hm D u mo rt . P iippo ( L ehm.) J . ( H ook.) f. N ee s H en ts c hel ( N ee s ) ( R ein w . ( N ee s ) ( Ca rr ing t on) J ö r g. ( L .) D u mo rt . H en ts c hel ( S p ru c e) ( H ook. ( S m.) ( N ee s ) ( L .) ( Sc h r e b .) ( H ook.) N ee s S t o le r ( R ein w . fl a gellife r Tab le ( Sc h r a d.) ( D i c k s .) ( Wa hlen b .) ( H ook. ( Ta y lo r ) ( G o tts c he) ( G r olle) ( N ee s ) w hi t eleggei c o r a llioide s G r a y H en ts c hel a s ellifo r mi ba r t a dend r oide s a tr o v i r en s o b tus ifoli u m w ood s ii a mphi b oli us di v a r i ca t pol y a n t ho s diploph y ll a m yr io ca r p us c on tr ac t a fl a vus flo t o v i a n us linden b e r gi a n us b i c us pid a t s c hi z ople ur a l a n c eol t ple ur a t di c r a n us v en tr i c o s a c oll a r i s p ru ino s a tr i c r en a t g r a v eolen s s ca l a r i Ba l a n t iop s i Ba r b ilopho z i a Ba zz a ni A del a n t h us B r e v i a n t h us C eph a lo z i C eph a lo z iell C h a e t oph y llop s i C hi a st o ca u lon C hilo s c y ph us C us pid a tu l D end r om a st igopho D in c kle r i a D iploph y ll u m E r emono tus H e t r o s c y ph us G eo ca l yx G o tts c heli a G y mnomi tr ion H e r b rtus Ha r p a n t h us J u nge r m a nni L eio c ole a L ep t o s c y ph us L io c hl a en L opho z i a Ma st igopho r a L epi c ole a Na r di a 128K. Feldberg, J. Vá≈ a, R.-L. Zhu & J. Heinrichs Acc .- N o. J 9849 3 5 Y 700000 Y 4 62303 Y 149855 Q 3 1 2 499 Q 9 003 11 Q 026 591 AY 60 8 036 AY 4 6232 1 AY 5 07 4 23 AY 4 6232 9 AY 4 6233 1 DQ 3 1 2 5 0 9 GQ 9 0030 4 GQ 9 00306 GQ 9 0033 4 GQ 9 003 15 G en Ba nk s eq u en c e A f r i ca R i ca Z e a l nd a ll SA fo r B r a z il N e w C hile in c l u ded a r e L] ( GOET ) C o st a 26 5 ]U r bc [ 111 07 [ r ef.no. ( GOET ) C hile ( JE ) P ol a nd s .n. CR -99-114 2 ( GOET ) G e r m a n y AY 6 9999 n u m b e rs ( GOET ) I ndone s i aA ( c on t in u ed) 151 V o u c he r O igin ( GOET ) N o rw a y DQ 3 1 2 4 7 5 V e rw imp ( GOET ) G e r m a n y GU 59 30 95 ( GOET ) A ustr i aD 4 33 9 CH - 00 7 1 ( GOET ) Ec u a do r GQ 9 0032 8 ( H 3 19 2330 ) Ta n z a ni aD & 20 9 3 5 ( HIRO ) Ja p a n GU 59 30 9 3 ( HIRO ) Ja p a n GU 59 30 94 ( HSNU ) C hin aG ( GOET ) G e r m a n y GU 59 30 9 6 ( GOET ) I ndone s i aF ( JE ) G e r m a n y GQ 9 00302 ( JE ) S o ut h ( H ) C hin aA ( H ) C hin aA 0 4 2 1 ( GOET ) M e x i c o DQ 3 1 2 51 0 ( GOET ) G e r m a n y AY 700003 4 70 1 4 3 91 K 4 ( H ) C hile a n l y e s . p.p. p.p. G en ba nk G r o t h Za r no w ie c ( GOET ) Ec u a do r GQ 9 003 5 7 a l. a l. a l. ( E ) N ep a l GQ 9 003 1 2 ( DUKE ) USA 449 3 B ry o 26/33 11 067 11 0 14 9 0066/ F & & 2 9 7 8 e t e t e t 5591 2 4 676 8 C r a nd ll- S t o le 5888 5 33 4 67 1 3/ 4 ML 159 6 1 Ja p a n AY 4 62327 262 2023 3 45 2 8499 F r a n z ing Sc h ä fe r - V e rw imp a l. & E 18 0 9 1 7022 & & e t RSA 2006072 8-14 a nd oponen h u ed rz eiko s eq u en c e . MP Sc h ä fe r - V e rw imp H ein r i c h s G r a d st ein S t o le r M i zut a ni H ein r i c h s H ein r i c h s O hni s hi O hni s hi Sc h r öde H ol z L ong A rts Sa u e r P ó c s G r a d st ein in us ed Ha t c he r H yv önen mole c u l a r e t fo r ) K oponen Sc h ust . G lenn y F u lfo r d G r olle E v a n s H yv onen IZ R . M a l. A . IK S t eph. a l. e t tru n ca tu m s pe c imen Ha tt . Sc h ä fe r - V e rw imp Ta y lo r ) Ta y lo r ) e t a l. a l. e t S . e t B u c h H ein r i s e t ) f. e t f. Ta y lo r ) D u mo rt . H o r n a l. Kaa l. H . Ta y lo r ) e t Ma ss a l. G r olle e t e t F eld b e r g f. G o tts c he) Sc hiffn. B e r gg . H en ts c hel Ta x on V o u c he r f. C . F eld b e r g S p ru c e H ol z e t ( H ook. ( H ook. S t eph. 1 : D u mo rt . H ein r i c h s F eld b e r g ( N ee s ) M i tt . B ur gh a r d t F eld b e r g ( Ma rt .) ( H u d s .) Lac .) P edinoph y ll u m ( N ee s ) ( H ook. ( R ein w . Ha tt .) ( L .) ( S t eph.) ( N ee s ) ( H ook. G r olle ( DC .) II III IV VJ ( Sc h r e b .) ( N ee s ) Tab le ( S . II (de t . ( L inden b . ( D i c k s .) ( G o tts c he) ( L ehm.) S t eph. ( Sa nde ( L .) su b tr ifid u m c y m ba life r a tu loide s den u d a tu m in t e rru p tu m in t e rru p tu m a r gill ac e us oppo s i tu m min utus o b v a tu m l a minige r a s plenioide q u inq eden t a h y a lin in v e rs a a nom l a utu mn li s a utu mn li s a utu mn li s a utu mn li s a utu mn li s nipponi ca c olo r a t nipponi ca nipponi ca c on r e t a p ur a s c en ru b r i ca u li s nemo r e a c urv ifoli a N o w elli a O don t o s c hi m a Pa r a s c hi st o hil P edinoph y ll u m P edinoph y ll u m P l a gio c hilion P l a gio c hil N ee s io c y ph us Sca p a ni Sc hiffne r i a Sc hi st o c hil a S phenolo b us S oleno st om a S yzy giell a S yzy giell a S yzy giell a S yzy giell a S yzy giell a S yzy giell a S yzy giell a S yzy giell a S yzy giell a S yzy giell a S yzy giell a S yzy giell a S yzy giell a T e tr ac y m ba liell a T r i a nd oph y ll u m T r i t om a W e ttst eini a The systematic position of Pedinophyllum (Marchantiophyta: Jungermanniales)129 consensus of four maximally parsimonious trees (length 1.672 steps, consistency index excluding uninformative characters 0,31) is depicted in Fig. 1. The two accessions of Pedinophyllum interruptum form a monophyletic lineage with a BS of 100. This lineage is nested in the robust Plagiochilaceae (BS 94) represented by Chiastocaulon, Dinckleria , Plagiochila and Plagiochilion. Plagiochilaceae are placed in a well supported sister-relationship (BS 94) with a clade including several representatives of Lophocoleaceae ( Chiloscyphus Corda, Heteroscyphus Schiffn., Leptoscyphus Mitt., Tetracymbaliella Grolle) and Brevianthus flavus (Grolle) J.J. Engel et R.M. Schust. of the monogeneric family Brevianthaceae. The Lophocoleaceae/Brevianthaceae clade achieves a BS of 77. The published rbcL sequence assigned to Pedinophyllum truncatum is nested in a lineage with two accessions of Syzygiella nipponica (S. Hatt.) Feldberg et al. with a BS of 99. Two sequences from Japanese specimens identified as P. truncatum (“Syzygiella autumnalis I and III”) are placed sister to a lineage with three sequences of Syzygiella autumnalis (I, IV, V). The ML analysis yielded a single tree with ln=-9764,2164. The ML topology (Fig. 2) resembles the MP strict consensus topology. Differences are seen only in clades without or with low BS.

DISCUSSION

The present study introduces new DNA vouchers of Pedinophyllum interruptum . The position of these accessions in the MP and ML trees (Figs 1, 2) lends strong support to the placement of Pedinophyllum in Plagiochilaceae. This position is supported by the “plagiochilid” perianths of Pedinophyllum , being laterally compressed and bilabiate, with the mouth truncate and not contracted. Pedinophyllum truncatum shares the perianth structure with the generitype (Gao, 2000, p. 262, plate 137) and thus also conforms with the Plagiochilaceae. The DNA voucher Koponen 46768 from China (GenBank no AY462321) was not available for study, however, its position in the robust Syzygiella nipponica clade lends support to the hypothesis of Crandall-Stotler et al. (2009) that the assignment to Pedinophyllum was based on a misidentification. Sterile material of the Syzygiella autumnalis / S. nipponica complex may easily be confused with Pedinophyllum , as is seen in the position of two further sequences generated from Japanese “ Pedinophyllum truncatum ” specimens that each include a mixture of various liverworts (HIRO, Ohnishi 2022 and 3452). These specimens include single perianth bearing shoots of Pedinophyllum and sterile Syzygiella shoots. DNA extraction from the Pedinophyllum material remained unsuccessful. However, we were able to sequence the sterile Syzygiella material (Figs 1, 2, “ S. autumnalis II, III). This material is nested in a robust lineage of Syzgyiella nipponica and S. autumnalis , and placed sister to S. autumnalis s.str. (MP tree, Fig. 1) or nested in the polytomous S. autumnalis clade (ML tree, Fig. 2). Fertile material of Jamesonielloideae can easily been separated from Plagiochilaceae by the plicate perianths narrowed towards their mouth. The rbcL sequence of Brevianthus flavus of He-Nygrén et al. (2006) places this species within Lophocoleaceae, sister to Tetracymbaliella . Engel & Schuster (1982) pointed out a striking resemblance of Brevianthus J.J. Engel et R.M. Schust. and Clasmatocolea Spruce [= Chiloscyphus (Hässel de Menéndez, 1996; Hentschel et al., 2007)] in the Lophocoleaceae. However, based on morphological peculiarities such as lack of underleaves and presence of androecia 130 K. Feldberg, J. Vá≈ a, R.-L. Zhu & J. Heinrichs

Fig. 1. Strict consensus of four most parsimonious trees with bootstrap percentage values > 50indicated at branches. Pedinophyllum interruptum is placed in the well-supported Plagiochilaceae. The systematic position of Pedinophyllum (Marchantiophyta: Jungermanniales)131

Fig. 2. Optimal tree (ln = -9764.2164) obtained in a maximum likelihood analysis under the GTR + G + I model. Bootstrap percentage values (BPV) > 50 are given at branches with PAUP*-BPV printed in regular letters, but GARLI-BPV in bold. 132 K. Feldberg, J. Vá≈ a, R.-L. Zhu & J. Heinrichs and perianths on branches lacking normal leaves, Engel & Schuster (1982) proposed a separate family, Brevianthaceae. Generic classification of Lophoco- leaceae is subject to controversy (e.g., Hentschel et al., 2007; Crandall-Stotler et al., 2009) and deserves further study. The present investigation strongly supports inclusion of Brevianthaceae in studies aiming at clarifying the global phylogeny of Lophocoleaceae. Acknowledgements. Financial support by the German Research Foundation (DFG grant HE 3584 / 4 to JH) is gratefully acknowledged. JV acknowledges support from the Ministry of Education of the Czech Republic through project No. 0021620828. RLZ appreciates support from the National Natural Science Foundation of China (NSFC, nos. 30825004, 30670155) and the Program of Shanghai Subject Chief Scientist (no. 08XD14016). We thank the director and curator of the Hiroshima University herbarium (HIRO) for the loan of specimens and the permission to extract DNA, and numerous colleagues for checking their herbaria for recently collected material of Pedinophyllum truncatum .

REFERENCES

AKAIKE H., 1974 — A new look at the statistical model identification. IEEE Transactions on Automatic Control 19: 716-723. CRANDALL-STOTLER B., STOTLER R.E. & LONG D.G., 2009 — Phylogeny and classification of the Marchantiophyta. Edinburgh journal of botany 66: 155-198. ENGEL J.J. & SCHUSTER R.M., 1982 — Austral Hepaticae XV. Brevianthaceae: A monotypic family endemic to Tasmania. The bryologist 85: 375-388. FELDBERG K., VÁ~ A J., LONG D.G., SHAW A.J., HENTSCHEL J. & HEINRICHS J., 2010 — A phylogeny of Adelanthaceae (Jungermanniales, Marchantiophyta) based on nuclear and chloroplast DNA markers, with comments on classification, cryptic speciation and biogeography. Molecular phylogenetics and evolution 55: 293-304. FELSENSTEIN J., 1985 — Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39: 783-791. GAO C., 2000 — Pedinophyllum (Lindb.) Lindb. In : Gao C. & Cao T. (Eds) Flora Yunnanica, vol. 17. Beijing, Science Press, pp. 261-263. GRADSTEIN S.R., WILSON R., ILKIU-BORGES A.L. & HEINRICHS J., 2006 — Phylogenetic relationships and neotenic evolution of Metzgeriopsis (Lejeuneaceae) based on chloroplast DNA sequences and morphology. Botanical journal of the Linnean society 151: 293-308. GROTH H. & HEINRICHS J., 2003 — Reinstatement of Chiastocaulon Carl based on evidence from nuclear ribosomal ITS and chloroplast gene rps 4 sequences. Plant biology 5: 615-622. HÄSSEL DE MENÉNDEZ G.G., 1996 — Reduction of Clasmatocolea Spruce and Xenocephalozia Schust. to the synonymy of Chiloscyphus Corda. Some rehabilitations and new combinations in Chiloscyphus (Hepatophyta). Nova Hedwigia 63: 493-516. HALL T.A., 1999 — BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic acid symposia series 41: 95-98. HE-NYGRÉN X., AHONEN I., JUSLÉN A., GLENNY D. & PIIPPO S., 2004 — Phylogeny of liverworts – beyond a leaf and a thallus. In : Goffinet B., Hollowell V.C. & Magill R.E. (Eds), Molecular systematics of bryophytes , Saint-Louis, Missouri Botanical Garden Press, pp. 87-118. HE-NYGRÉN X., JUSLÉN A., AHONEN I., GLENNY D. & PIIPPO S., 2006 — Illuminating the evolutionary history of liverworts (Marchantiophyta) – towards a natural classfication. Cladistics 22: 1-31. HEINRICHS J., GRADSTEIN S.R., WILSON R. & SCHNEIDER H., 2005 — Towards a natural classification of liverworts based on the chloroplast gene rbcL. Cryptogamie, Bryologie 26: 131-150. HEINRICHS J., LINDNER M., GROTH H., HENTSCHEL J., FELDBERG K., RENKER C., ENGEL J.J., VON KONRAT M., LONG D.G. & SCHNEIDER H., 2006 — Goodbye or welcome Gondwana? – insights into the phylogenetic biogeography of the leafy liverwort Plagiochila with a description of Proskauera , gen. nov. (Plagiochilaceae, Jungermanniales). Plant systematics and evolution 258: 227-250. The systematic position of Pedinophyllum (Marchantiophyta: Jungermanniales)133

HENTSCHEL J., WILSON R., BURGHARDT M., ZÜNDORF H.-J., SCHNEIDER H. & HEINRICHS J., 2006 — Reinstatement of Lophocoleaceae () based on chloroplast gene rbcL data. Plant systematics and evolution 258: 227-250. HENTSCHEL J., FELDBERG K., ZÜNDORF H.-J., HELLWIG F.H., SCHNEIDER H. & HEINRICHS J., 2007 — The systematic position of Pachyglossa and Clasmatocolea (Jungermanniopsida: Lophocoleaceae) inferred from nrDNA ITS sequences and morphology. Taxon 56: 1136-1142. HILLIS D.M. & BULL J.J., 1993 — An empirical test of bootstrapping as a method for assessing the confidence in phylogenetic analysis. Systematic biology 42: 182-192. INOUE H., 1958 — The family Plagiochilaceae of Japan and Formosa. I. Journal of the Hattori botanical laboratory 19: 25-59. POSADA D. & CRANDALL K.A., 1998 — Modeltest: testing the model of DNA substitution. Bioinformatics 14: 817–818. SWOFFORD D.L., 2003 — PAUP*, phylogenetic analyses using parsimony (*and other methods), version 4.0b10. Sinauer, Sunderland, Mass. TAVARÉ S., 1986 — Some probalistic and statistical problems in the analysis of DNA sequences. Lectures on mathematics in the life sciences 17: 57-86. ZWICKL D.J., 2008 — GARLI (Genetic Algorithm for Rapid Likelihood Inference). Version 0.96, available from: .