Newton2009chap12.Pdf
Total Page:16
File Type:pdf, Size:1020Kb
Mosses (Bryophyta) Angela E. Newtona,*, Niklas Wikströmb, and a single spore capsule and oJ en with a complicated dehis- A. Jonathan Shawc cence (dispersal) mechanism (peristome). Spores are aDepartment of Botany, Natural History Museum, London SW7 5BD, formed by meiosis and usually dispersed as monads. UK; bDepartment of Systematic Botany, Evolutionary Biology Centre, 7 e taxonomic diversity of living mosses reP ects the Uppsala University, Norbyvägen 180 75236, Uppsala, Sweden; morphological diversity, and is arranged in a pectin- cDepartment of Biology, Duke University, Durham, NC, 27708 USA ate grade of several small or very small but highly dis- *To whom correspondence should be addressed (a.newton@nhm. ac.uk) tinct clades, a few rather larger but also distinct clades (e.g., Sphagnales, Polytrichales) and two very large and poorly resolved clades, the Dicranidae and Bryidae, that Abstract together account for 90% of extant moss species diversity. 7 e basal grade, the Dicranidae and part of the Bryidae Living mosses (ca. 13,000 sp.) constitute the Phylum are acrocarpous, with the principal vegetative axis Bryophyta, with eight classes divided into acrocarpous terminated by gametangia, and consequently with ter- mosses (23 orders), not an evolutionary group, and pleuro- minal formation of sporophytes. A monophyletic group carpous mosses (4–7 orders, 42% of extant species). Two sub- within the Bryidae is pleurocarpous, with gametangia classes, Dicranidae (acrocarpous haplolepidae) and Bryidae (diplolepidous- alternate mosses, with both acrocarpous and pleurocarpous members), account for 90% of extant species. The moss timetree shows lineage origin at ~380 million years (Ma) ago, the split between Haplolepidae and Diplolepidae at 220–195 Ma, and appearance of the fi rst pleurocarp lineages at ~173 Ma. Major diversifi cation occurred in the Cretaceous, 140–90 Ma. Mosses are photosynthetic plants that exhibit a wide range of morphologies, based on the life cycle of alterna- tion of haploid and diploid generations. 7 e gametophyte generation starts with haploid spores that develop into threadlike protonema, from which grow erect or creep- ing axes usually up to a few centimeters tall. 7 e plants are oJ en branched, with leaves that are mostly one cell thick and usually radially arranged (Fig. 1). 7 ey form small cushions, velvety swards, open turfs, tuJ s, or deep mats. Long-lived clonal growth and specialized vegeta- tive reproduction are common and widespread, with sex- ual reproduction rare or unknown in some taxa. Plants are male or female, or bisexual, with various diB erent arrangements of gametangia. Male and female gametan- gia are distinct, with a basic morphology common to all mosses (and to most of the early-diverging land-plant lin- eages) but with variation in some groups. Motile sperm- atozoa travel though surface water to fertilize sedentary eggs. 7 e resulting diploid embryo grows into a sporo- Fig. 1 Wall screw-moss (Tortula muralis) from a brick wall in phyte that is largely dependent on the gametophyte, with Richmond, England. Credit: A. E. Newton. A. E. Newton, N. Wikstrom, and A. J. Shaw. Mosses (Bryophyta). Pp. 138–145 in e Timetree of Life, S. B. Hedges and S. Kumar, Eds. (Oxford University Press, 2009). HHedges.indbedges.indb 113838 11/28/2009/28/2009 11:25:48:25:48 PPMM Eukaryota; Viridiplantae; Bryophyta 139 Plagiotheciaceae Meteoriaceae Fontinalaceae Cryphaeaceae Pterobryaceae Thuidiaceae 26 Entodontaceae Hypnaceae 16 Brachytheciaceae 28 Hylocomiaceae Leucodontaceae Neckeraceae Hookeriales/Hypnales Hypnaceae Stereophyllaceae Fabroniaceae 15 Lepyrodontaceae Catagoniaceae Bryidae Leucomiaceae 29 27 Pilotrichaceae 13 Hookeriaceae 23 Trachylomataceae Rutenbergiaceae Hypopterygiaceae Ptychomniaceae (Ptychomniales) Aulacomniaceae (Aulacomniales) 11 Pterobryellaceae 25 Braithwaiteaceae 21 Racopilaceae Hypnodendraceae Hypnodendrales Hypnodendrales Orthodontiaceae (Orthodontiales) 8 Rhizogoniaceae (Rhizogoniales) (continued on next page) D C P Tr J K Pg Ng PALEOZOIC MESOZOIC CZ 300 200 100 0 Million years ago Fig. 2 Continues terminating specialized lateral branches, so that sporo- A similar morphological transition is seen in the struc- phytes develop on branches. 7 is innovation has been ture of the dehiscence mechanism of the sporophyte. suggested to be a key in the evolution of the branching 7 e earliest diverging lineages mostly have simple linear structure, contributing to the enormous species diversity dehiscence while later diverging lineages have circumscis- in this group (1). sile dehiscence and (usually) a peristome. 7 e peristome HHedges.indbedges.indb 113939 11/28/2009/28/2009 11:25:52:25:52 PPMM 140 THE TIMETREE OF LIFE (continued from previous page) Hedwigiaceae (Hedwigiales) Bartramiaceae (Bartramiales) Mniaceae Bryaceae Bryales Bryales Orthotrichaceae (Orthotrichales) Bryidae 10 Splachnaceae 19 Meesiaceae 6 Timmiaceae (Timmiales) Splachnales 17 Scouleriaceae (Scouleriales) Pottiaceae (Pottiales) 20 Ditrichaceae 7 14 Dicranaceae 18 Fissidentaceae 5 Dicranales Schistostegaceae 22 12 Seligeriaceae (Seligeriales) Dicranidae Ptychomitriaceae 24 Grimmiaceae 4 Funariaceae (Funariales) Grimmiales 9 Encalyptaceae (Encalyptales) Diphysciaceae (Diphysciales) Buxbaumiaceae (Buxbaumiales) 3 Tetraphidaceae (Tetraphidales) Polytrichaceae (Polytrichales) 2 Oedipodiaceae (Oedipodiales) Andreaeaceae (Andreaeales) 1 Andreaeobryaceae (Andreaeobryales) Takakiaceae (Takakiales) Sphagnaceae (Sphagnales) D C P Tr J K Pg Ng PALEOZOIC MESOZOIC CZ 300 200 100 0 Million years ago Fig. 2 A timetree of Bryophyta. Classifi cation follows Bell et al. (35) for the pleurocarpids and Goffi net and Buck (4) for the remaining taxa. Abbreviations: C (Carboniferous), CZ (Cenozoic), D (Devonian), J (Jurassic), K (Cretaceous), Ng (Neogene), P (Permian), Pg (Paleogene), and Tr (Triassic). itself shows a transition from massive teeth composed walls between three concentric rings of cells. 7 e patterns of multiple cells (nematodontous) to more delicate and of the cell wall remnants on the peristome surfaces have P exible teeth composed of cell walls (arthrodontous). In allowed the correspondence of the structures to be iden- the arthrodontous mosses the peristome is composed of tiA ed (2). Mosses with both peristome rings are termed one or two rings of structures, the outer exostome teeth diplolepidous, and have an exostome that is robust and and the inner endostome segments, derived from the cell P exes with changes in humidity, while the endostome is HHedges.indbedges.indb 114040 11/28/2009/28/2009 11:25:52:25:52 PPMM Eukaryota; Viridiplantae; Bryophyta 141 Table 1. Divergence times and their confi dence/ the peristome is known in many diB erent taxa. Both the credibility intervals (CI) among Bryophyta, based haplolepidous- and diplolepidous-alternate forms appear on ref. (1). to be derived from the diplolepidous-opposite group, but Timetree this area of the topology is not yet strongly resolved or Node Time CI supported. However, these forms comprise very large monophyletic groups, the haplolepidous Dicranidae and 1379.0400–362 the diplolepidous-alternate Bryidae (sensu 4), with 30% 2329.0342–304 and 60%, respectively, of extant species diversity (1). 3317.0334–295 ClassiA cation of the mosses has P uctuated wildly over 4292.0316–280 the centuries since the starting-point publication by 5246.0272–230 Hedwig in 1801 (5), depending on whether the sporophyte 6219.0243–205 or gametophyte generation was given precedence (6). 7203.0220–176 Certain groups in which the peristome is highly reduced 8195.0216–181 or absent have been particularly problematic. However, some degree of stability was introduced with the work of 9187.0209–162 Brotherus (7) and Fleischer (8) in the early twentieth cen- 10 184.0 204–165 tury, although numerous small and not-so-small changes 11 173.0 194–161 have continued to be made in taxon circumscription and 12 156.0 188–144 relationships at all levels, and opinion has diB ered on 13 151.0 173–141 particular taxa. For example, the Polytrichales, which 14 149.0 175–130 have relatively well-developed vascular tissue, have been 15 143.0 165–131 placed near the beginning (9) or end (8) of classiA ca- 16 141.0 157–123 tions, with implications of a primitive or derived status, 17 136.0 159–111 respectively. 7 e relatively small size and simple struc- tures of mosses appears to have led to extensive parallel- 18 121.0 145–101 ism and convergence, making the use of morphological 19 116.0 138–95 characters for classiA cation particularly di1 cult. 20 111.0 141–96 Cladistic methodology was adopted early by bryolo- 21 111.0 124–88 gists, one of the earliest applications of Hennigian prin- 22 109.0 134–88 ciples was a generic revision (10) of the moss Family 23 107.0 136–93 Mniaceae in 1968, and in 1984 a morphological cladistic 24 105.0 131–82 analysis (11) of the bryophytes established the very basic 25 88.0 102–67 elements of the pectinate grade (Sphagnales (Andreaeales (Tetraphidales (Polytrichales (Buxbaumiales (Bryales)))))) 26 71.0 96–56 most of which is still accepted. A pioneering series of cla- 27 71.0 92–61 distic analyses (using morphological data) explored the 28 67.0 118–54 relationships of the pleurocarpous mosses (12). However, 29 47.0 61–39 it was not until the advent of DNA sequencing that su1 - cient data were available to explore relationships in detail. Since the late 1990s studies using single plastid or mul- more delicate but less P exible. Changes in positions and tigene phylogenies (oJ en including