J Hattori Bot. lab. No. 84: 37-47 (July 1998)

THE ARCHITECTURE OF THE (MUSCI)

R. S. TANGNEY1

ABSTRACT. An analysis of the architecture of the Lembophyllaceae is presented and its value in the of the family is assessed. The Lembophyllaceae produce a variety of growth forms, rang­ ing from creeping mats to loose erect-ascendant wefts and pendant fronds. Characters of stem orien­ tation, branching, rhizoid distribution, leaf orientation, size and shape, and the production of determi­ nate or indeterminate growth are used to describe merophyte development. Variation in the pattern of merophyte development yields an architectural pattern common to taxa, and the genera differ from each other in their relative expression of this pattern. The same architecture is therefore seen to un­ derlie the various growth forms exhibited.

K EY WORDS: , Lembophyllaceae, architecture, ontogeny, heterochrony.

INTRODUCTION The Lembophyllaceae are a Southern Hemisphere family of pleurocarpous mosses, occurring in Australasia and southern South America. During its history, the family has in­ cluded a diversity of unrelated elements that in recent years have been transferred to other families (Buck 1980, 1994; Vitt 1984; Crum 1991 ; Tangney 1996, I 997b ). While the fami­ ly recognised in a more restricted sense (than that, for example, of Brotherus 1924- 25) is morphologically more coherent, there remain difficulties in the separation of the genera. These problems are related to the extreme variability of species in the family. This plastici­ ty is associated with variation in the underlying architecture (Tangney 1996). This descrip­ tion of the architecture aims to provide a basis for the understanding of the variability of the species, as well as the inter-relations of the genera.

ARCHITECTURE AND GROWTH FORM. The term architecture is not generally used in reference to bryophytes. Rather, the cat­ egories of growth-form and life-form have traditionally been employed to describe the morphological aspects of branching pattern and overall appearance (physiognomy) of bryophytes, respectively. While there is a clear difference in the definition of these terms, their usage has been ambiguous and the value of growth-form in morphological analysis has been eroded. Magdefrau (1982) considered growth-forms to be the" ... genetically fixed method of ramification'', i.e., characteristic of an individual, and life-forms to be an " ... as­ semblage of individuals and growth-form, modified by external conditions'', i.e., character­ istic of a group of individuals. Growth-form is therefore subordinate to life form as it is a component of the latter. While the distinction between the two categories is clear, the use of the terms has been confusing. For example, Magdefrau (1982) cited earlier authors' defi­ nitions of growth form as being strictly morphological, yet considered growth form to be modified by the external environment. 1 Department of Botany, University ofOtago, P 0 Box 56, Dunedin, New Zealand. 38 J. Hattori Bot. Lab. No. 84 I 9 9 8

Gimingham and Robertson (1947) produced a growth-form classification based on that of Meusel (1935). Their modifications to Meusel's detailed system explicitly gave more importance to "general growth morphology" than to branching pattern. Thus their 'growth form' became an index of environmental influence rather than genetically fixed. Therefore, both life-form and growth-form systems have come to produce very simi­ lar classifications, even sharing descriptive terms, for example turf, weft, cushion and mat (see Magdefrau 1982 and Richards 1984). This, coupled with the environmental influence associated with growth-forms, has led to a perception that characters of the growth-form have little value in taxonomy. Because of the ambiguity surrounding growth-forms, the term architecture, as applied to vascular , is preferred. Both Mishler and De Luna ( 1991) and La Farge-England (1996) have shifted the emphasis of growth-form study back to an architectural analysis. The architecture of a is the result of meristematic activity, a developmental sequence which is independent of physiognomy, biological type or taxonomic position (Halle and Oldeman 1975). The use of the term architecture emphasises the analysis of the develop­ mental branching pattern rather than overall appearance (Mishler and De Luna 1991 ). Bryophytes may therefore exhibit similar architectures to other groups of plants, and differ­ ent 'growth-forms' may be produced by similar architectures. Mishler and De Luna ( 1991) outlined a hierarchical framework for ontogenetic de­ scription ofbryophytes. It utilises some terminology applied to vascular plants and consists of five hierarchical levels of development; the cell, the metamer (=the merophyte), the module (=the branch), the branch system (=the shoot), and the shoot system. Differentia­ tion occurs at each level and may vary within each level depending on the developmental level of the higher stage. For example, merophyte development may vary both within a branch and between branches of the same shoot. The pattern of merophyte development and the timing and pattern of the structures produced, therefore yields an architectural pat­ tern that may be used to compare taxa.

THE LEMBOPHYLLACEAE The Lembophyllaceae are characterised by morphological variation within taxa such that no characters are discontinuous between the genera. Dixon ( 1927) considered that the species were easy enough to recognise, but that the genera were not. For example, Camp­ tochaete exhibits frequent variation on its dendroid-stipitate architecture. This flexibility of form, combined with vigorous growth produces plants that blur species boundaries. Some of these forms (called 'deflexa' forms by Dixon, 1927) have been a source of superfluous names, and previous authors have noted taxonomic difficulties associated with variability (Dixon 1927, Sainsbury 1955, Scott and Stone 1976, Crum 1991 ). This variability is a function of plant architecture (Tangney 1996), and this explication of the architecture and variation of the Lembophyllaceae is aimed at an understanding of the variation observed. In pleurocarpous mosses, growth of the apical cell is not terminated by gametangial production, i.e. module growth is not determinate as in acrocarpous mosses. Therefore, there is the potential for greater differentiation within modules. In this analysis, characters are utilised as indicators of merophyte development within the levels of module and branch R. S. TANGNEY: Architecture of the Lembophyllaceae 39 system (shoot). This pattern ofmerophyte development is used to compare the taxa. Characters utilised: Stem orientation Orthotropy: growth mostly erect with radial symmetry Plagiotropy: growth mostly horizontal with flattened (complanate) growth Branching (presence or absence) Growth Monopodial: the continuous activity of a single apical cell. Sympodial: growth from lateral apical cells. For example, when growth of the frond axis api­ cal cell stops, growth of new modules, comes from lateral apical cells. Growth Determinate: Growth producing a fixed structure or number of structures only, e.g. a leaf, a sporophyte, or a branch of fixed length. Indeterminate: Growth producing a succession of structures of no fixed number, e.g., the main axis of a pleurocarpous . Rhizoid distribution Leaf orientation, size and shape

DESCRIPTION OF TAXA exhibits a diversity of architectural pattern which encompasses that of the other genera. Camptochaete is described in detail, and the other genera are compared to it. The genera are seen to differ from each other in the relative expression of the same un­ derlying pattern. Observations are based on studies of herbarium, field, and cultivated ma­ terial. Camptochaete Reichdt. Ten species in two sections (Tangney l 997a). Details of the archi­ tecture of the two sections differ. The architecture of species in Sect. Camptochaete is de­ scribed, and differences with sect. Thamniella are noted below.

Camptochaete sect. Camptochaete Growth form (Fig. 1.). Plants are erect-ascendant, forming loose wefts, and occasion­ ally, distal stems and branches may be pendant. Rarely, plants are found in unattached loose-lying "moss-balls" on the forest floor. Stem differentiation present. Erect stems are produced from creeping stolons. Stems consist of a lower unbranched stipe and an upper branched frond. Stolon growth is pla­ giotropic; creeping with rhizoids scattered. Leaves erect and imbricate, often erose. Stipe growth is orthotropic, developed from the creeping stolon by the stolon tip becoming erect. Leaves of the stipe are erect and imbricate. The stipe is usually unbranched and rhizoids are absent. Fronds are plagiotropic; with irregular bi-tripinnate branching usually produc­ ing flattened fronds. Fronds are determinate, with lateral branches determinate and the frond axis also usually determinate. Reiteration. Reiteration of the branch system occurs either sympodially or monopodi­ ally. Sympodial innovations arise as lateral branches from the base of the stipe, and are ini­ tially plagiotropic (stolon), become orthotropic (stipe), and then plagiotropic (frond). 40 J. Hattori Bot. Lab. No. 84 I 9 9 8

Camptochaete

a

Fig. I. Architecture of Camptochaete. a. stolon, b. stipe, c. frond, d. stoloniferous or flagelliferous growth of the frond, e. elongate or 'deflexa' form. Growth of module a-b-c is monopodial. Reiteration is either sympodial or monopodial at d., or sympodial at base of the stipe (b). Typically Camptochaete is stipitate (a-b-c) and exhibits the full range of architec­ tural variation. Arrows indicate the positions where new modules may be produced. See text for details.

Monopodial reiteration of the branch system occurs through the frond axis having indeter­ minate growth. In this case frond development (lateral branching) is suppressed, leaf orien­ tation changes from erect-spreading to erect, and stem orientation becomes positively geot­ ropic. The tips become rhizoidal and root. This stoloniferous tip is initially plagiotropic and becomes orthotropic to produce a new branch system of stipe and (plagiotropic) frond. In this way an arching and rooting architecture is produced (Tangney 1997a). New branch systems may also be produced sympodially from the frond as dominant lateral branches of indeterminate growth. These innovations are characterised by having erect-appressed leaves at the base. Sometimes the frond axis continues growth to produce elongate or 'deflexa' (Dixon 1927) forms. These generally trailing or pendant fronds are characterised by a stoloniferous appearance and a tendency to produce erect-appressed leaves, distant branches and scattered rhizoids. Here there is a contrast between the frond axis and the branches with respect to leaf orientation.

Camptochaete sect. Thamniella Camptochaete sect. Thamniella is distinguished vegetatively from C. sect. Camp­ tochaete by having patent and distant stipe leaves, rather than erect-appressed and imbri­ cate. Otherwise it exhibits the same architectural pattern. When reiteration from the frond R. S. TANGNEY: Architecture of the Lembophyllaceae 41

Lembophyllum

d ~

Fig. 2. Architecture of Lembophyllum. a. stolon, b. stipe, c. frond. Growth of module a-b-c is monopodial. Reiteration is either sympodial or monopodial at d., or sympodial at the base of the stipe (b ). Typically Lembophyllum exhibits frond development with less stolon and stipe. Arrows indicate the positions where new modules may be produced. See text for details. occurs in sect. Thamniella, there is a similar repetition of stipe leaf characters in sect. Thamniella . Fifea Crum. The monotypic Fifea exhibits the same architectural pattern as Camptochaete sect. Camptochaete, and is not described separately. Lembophyllum Lindb. Three species in two sections (Tangney l 997b ). Growth form (Fig. 2.). Species of Lembophyllum are generally smaller plants than the more robust members of Camptochaete. Species of lembophyllum commonly produce wefts or cushions. Stem differentiation is present. Erect stems are produced from somewhat abbreviated stolons. Stems consist of a lower unbranched orthotropic stipe and an upper branched pla­ giotropic frond. Typically the branches of the frond are longer than in Camptochaete and the fronds are usually pinnate rather than bi- tri-pinnate as in Camptochaete. The stipes are also less well developed in Lembophyllum, which is perhaps a function of plant size. There is differentiation of leaves between the stipes and fronds. Leaves of the stipes are smaller, less concave, more erect and imbricate to distant. Therefore the stipes are narrower than the frond axis and frond branches. Reiteration. New stems are produced sympodially as basal innovations of stipes, or as strongly oriented dorsal innovations of the frond. Typically distal frond axes or branch tips may lose determinacy and produce reiterations monopodially or sympodially respectively. Branching is suppressed and leaves become smaller and more erect imbricate to produce stoloniferous tips, as in Camptochaete. Fallaciella Crum. One species. Growth form (Fig. 3.). Plants are small. They characteristically form closely appressed mats or low dense wefts. Stem differentiation is present. Erect stems are produced from stolons. Lower un- 42 J. Hattori Bot. Lab. No. 84 I 9 9 8

F allaciella

e a

Fig. 3. Architecture of Fallaciella. a. stolon, b. stipe, c. frond, d. stoloniferous or fla­ gelliferous growth of the frond, e. elongate or defiexa forms. Growth of module a-b-c is monopodial. Reiteration is either sympodial or monopodial at d., or sympodial or monopo­ dial at the stipe base (b) or stolon (a). Typically Fallaciella exhibits frond development with weak stolon and stipe, but often produces elongate forms. See text for details. branched stipes are sometimes produced. Typically plants consist of erect-ascendant to prostrate stems with dorsally oriented branches. Fronds are bi- tri-pinnate. Reiteration. Reiteration of the stems is by basal innovations of the stipe (sympodial), lateral innovation of the stolon or by distal prolongations of the frond, sympodially or monopodially as in Camptochaete. Elongate forms are often produced from the fronds, typically with an axis exhibiting leaf differentiation and distant lateral branches. These elongate forms are similar to those produced in Camptochaete. Broth. Two species in two sections (Tangney 1997b ). The two species differ in their architecture and are described separately.

Weymouthia cochlearifolia Growth form (Fig. 4.) . Weymouthia cochlearifolia is a relatively robust plant, usually forming dense wefts or turfs. It is sometimes pendant. Stem differentiation present. Erect stems are produced from creeping stolons. Erect stems are determinate and simple to sparsely branched. There is weak differentiation of leaves between creeping stems and erect branches. More robust erect stems may produce stipes with differentiated leaves. These stipitate shoots may arise from stolons that become erect and grow orthotropically, or as lateral branches of the creeping axis. Reiteration. Growth of the erect stems is usually as lateral innovations of the creeping stem, and thus monopodial. Occasionally tips of the erect stems become stoloniferous and rhizoidal, and then become erect again to reiterate the branch system. Branched flagellifer­ ous shoots may be produced at branch tips. Elongate monopodial forms may also become pendant. In such instances, there is weak differentiation of stem and branch leaves and branches are distant. R. S. TANGNEY: Architecture of the Lembophyllaceae 43

Weymouthia cochlearifolia

Fig. 4. Architecture of Weymouthia cochlearifolia a. stolon, b. stipe, c. frond, d. stoloniferous or flagelliferous growth of the frond, e. elongate or deflexa form. Growth of the module a-b-c is monopodial. Reiteration is either sympodial or monopodial at d., or sympodial at the base of the stipe (b ). Typically Wey­ mouthia cochlearifolia exhibits monopodial growth of a creeping axis with erect lateral branches which is often pendant (e). Stipitate growth a-b-c is less common. Arrows indicate the positions where new modules may be produced. See text for details. Weymouthia-- mollis

Fig. 5. Architecture of Weymouthia mollis. e. elongate frond. Growth of the frond is monopodial with lateral branches produced from the central axis. Reiteration is either monopodial or sympodial. Stipi­ tate growth is absent. See text for details. 44 J. Hattori Bot. Lab. No. 84 I 9 9 8

Weymouthia mollis Growth form (Fig. 5.). Weymouthia mollis typically forms extensive pendant fronds from twigs and branches of trees and shrubs. Stem differentiation is present. Short creeping stems give rise to pendant monopodial shoots of determinate growth. There is leaf differentiation between the main axis and later­ al branches. Leaves of the main axis are erect imbricate, whereas those of the branches are erect-spreading. Stem leaves are also longer and narrower. Weymouthia mollis does not pro­ duce stipitate forms, but differentiation in stem and branch leaves is similar to that of the elongate forms produced by the other taxa. Reiteration. New shoots arise as lateral branches of the creeping stem. The monopodi­ al units are determinate and reiteration of these is sympodial.

SUMMARY OF THE ARCHITECTURE The architectural pattern of the Lembophyllaceae is essentially dendroid-stipitate. The basic unit of reiteration, comprising stolon, stipe, and frond, is monopodial, with the central axis being the product of merophytes produced by a single apical cell. Merophytes in this unit are not all equal and vary in their differentiation. There are clear differences in phases of growth. Stolons, stipes, and fronds have different growth orientation and potential, branching and leaf size, shape, and orientation. Reiteration of the pattern involves repeti­ tion of earlier growth phases of the unit. For example, new innovations produced sympodi­ ally from basal regions of the stipe grow initially as stolon/stipe and then produce fronds. Similarly, new innovations produced sympodially from the frond are initially stipe-like, with erect appressed leaves at the base. When frond growth becomes elongate, stolons characters are introduced and stems are produced that become rhizoidal and creep over the substrate, often exhibiting features of the stolon and the frond mixed together. Therefore, these elongate forms represent reitera­ tion complexes, in which the orderly production of new innovations breaks down to pro­ duce diffuse growth of unsettled morphology. This architectural pattern finds its most complete expression in Camptochaete. Camp­ tochaete is typically dendroid stipitate and normally produces elongate forms. lembophyl­ lum, being typically frondose, produces both stipitate and elongate forms less often than Camptochaete. However, branches frequently end in stoloniferous and rhizoidal tips. Fall­ ciella also consists mainly of frond and often has elongate forms. Typically, it does not pro­ duce stipitate forms, and then these are usually weakly developed. As with lembophyllum, stipitate forms are usually larger plants. Weymouthia cochlearifolia usually produces monopodial forms, with densely crowded erect lateral stems arising from a creeping axis. The erect stems are not usually stipitate, but may be, and the branch tips can be stolonifer­ ous and rhizoidal. Loosely attached or pendant forms characteristically show leaf differen­ tiation. The monopodial axis can have leaves which grade from the typical characters of the frond axis leaves to characters typical of the stolon leaves. These latter stolon characters contrast strongly with the leaf characters of the branches. Weymouthia mollis differs from the other members of the family in not having stipitate forms or stoloniferous branch tips. The plants are normally pendant from short creeping stems. The pendant fronds are uni­ form and monopodial or sympodial. Lateral branches arise distantly and regularly. Whereas R. S. TANGNEY: Architecture of the Lembophyllaceae 45 in Weymouthia cochlearifolia the stolon characters of the creeping axis are variably pro­ duced, in Weymouthia mollis the leaf differentiation between the axis and branches is con­ stant. The pendant fronds of Weymouthia mollis are interpreted as equivalent to the elon­ gate reiteration complexes of the other taxa, but in this species the morphology is stable with little variability.

Phenotypic variability In Camptochaete, the specific variation previously noted can be related to architectur­ al variation. Individual species have a morphological 'core' of leaf characters associated with frond structure, and characters of this part of the plant, particularly those of the frond axis, are most useful in separating the species. Atypical material is associated with reitera­ tion of the basic architectural unit Stoloniferous prolongations of the frond and elongate monopodial forms produce leaf sizes and shapes that converge with those of other species. These morphological extremes formed by 'reiteration complexes' account for blurring of species boundaries. Similarly, as the genera share the underlying architectural pattern, generic boundaries are blurred by architectural variation. Differences amongst the genera are broadly de­ scribeable in terms of growth form/overall appearance, and the genera usually exhibit a part of the architectural pattern characteristic of the genus. Variation from typical material causes taxonomic problems. For example, elongate monopodial forms of Camptochaete ar­ buscula are difficult to separate from typical Weymouthia cochlearifolia, and stipitate forms of Weymouthia cochlearifolia are similar to typical Camptochaete arbuscu/a. Weymouthia cochlearifo/ia is most readily separated from by its strongly porose lamina! cells. Camptochaete arbuscula has lamina! cells not or only weakly porose.

Relations of the Genera The presence of the same architectural pattern in all genera of Lembophyllaceae sup­ ports the suggested relation of these genera (Tangney l 997b ). Weymouthia mollis is the only species not to exhibit the full range of architectural variation. The key components of this architectural pattern are considered as developmental homologues. The genera are in­ terpreted as the evolutionary result of heterochronic processes acting on an underlying ar­ chitectural pattern. Whether this pattern was fixed in the ancestral taxa, with the modern taxa evolving to exhibit different aspects of the pattern in their typical forms, or whether the modern taxa evolved out of ancestral taxa in which the morphology was not fixed (i.e. consisted of 'reiteration complexes') is not certain, and is the basis of ongoing phylogenet­ ic studies. Differences between the genera with respect to architecture are not clearcut and probably represents the influence of indeterminate growth in the gametophyte. Tan et al. ( 1996) suggested that Fallaciella gracilis was related to Camptochaete as it shared aspects of branching pattern with the latter, and that it was neotenous in origin, from an ancestral form similar to Camptochaete. This suggested heterochronic origin is support­ ed here. Whether neoteny is the heterochronic mode is unclear. 46 J. Hattori Bot. Lab. No. 84 I 9 9 8

Architecture and Growth Form Architectural analysis may be thought of as a search for repeatable units of plant con­ struction (Halle and Oldeman 1975) or modularity (modules). Mishler and De Luna (1991) outlined an ontogenetic scheme for mosses in which the module, as repeatable unit, was defined as the product of a single apical cell. Sympodial reiteration of this module gave rise to the higher level branch-system, which makes up "a single leafy shoot." In the architec­ tural pattern described here the axis of the stolon-stipe-frond is the product of a single api­ cal cell and therefore corresponds to Mishler and De Luna's module, but the leafy shoot of which it is the axis has lateral branches produced from lateral apical cells. Accordingly, the basic repeatable unit of construction in the Lembophyllaceae, or module in architectural terms, corresponds to Mishler and De Luna's branch system. As pleurocarpy produces plants in which the 'modules' are not all equal, their units of reiteration occur at a different hierarchical level than in acrocarps. There are few studies of architecture in mosses {for a detailed review, see Mishler and De Luna, 1991 .) Most have centred on the problem of acrocarpy-pleurocarpy-cladocarpy, or on overall orientation of growth, erect (orthotropy) vs creeping (plagiotropy), e.g., La Farge-England ( 1996), who grouped taxa into growth-forms emphasising perichaetial posi­ tion (acrocarpy, pleurocarpy or cladocarpy), nature of branching (monopodial or sympodi­ al), and direction of growth (orthotropy or plagiotropy). In the Lembophyllaceae, the growth-form in these terms is very variable. It is pleurocarpous, but branching and orienta­ tion of axes is various. The basic unit of reiteration ('module') is monopodial. It is initially plagiotropic, becomes orthotropic and then plagiotropic again. Reiteration is either sympo­ dial or monopodial. This pattern is very similar to that described for some Pterobryaceae (Argent 1973).

CONCLUSIO 'S The emphasis of architectural analysis is on developmental branching. The detailed analysis of the developmental pattern yields an understanding of both the morphology of species and the inter-relations species and genera. Unrelated taxa may share a particular ar­ chitectural pattern, but the comparison of related taxa is a useful source of evolutionary in­ formation. In the Lembophyllaceae, study of the architecture has: 1. Clarified morphological variation in species and the genera. 2. Provided an understanding of the basis of the 'growth-forms', and showed that, for example, mats, wefts and pendant forms have inter-related morphologies, i.e., they share the same architectural pattern. 3. Supported the evolutionary origin of the taxa by heterochrony 4. Provided the basis for ongoing developmental and phylogenetic studies on spore germination, heteroblasty, phase changes, and heterochrony.

ACKNOWLEDGEMENTS I thank the following for useful djscussions, Allan Fife, Michael Heads, Angela New­ ton, George Scott and Bastow Wilson. Rod Seppelt and Norton Miller both made helpful R. S. TANGNEY: Architecture of the Lembophyllaceae 47 comments on the manuscript. Support from the Science Research Committee of the New Zealand Lottery Grants Board is gratefully acknowledged.

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