Lembophyllaceae

Belongs within: Hypnales.

The Lembophyllaceae are a family of mostly epiphytic mosses found predominantly in the Southern Hemisphere, species of which generally have a weft-forming or pendent growth form. A stipitate-frondose growth habit, with leaves that differ between stipes, frond axes and branches, is found in members of the genus Camptochaete.

The trials and tribulations of tree moss

Published 17 May 2011

The Lembophyllaceae are a family of mosses found most abundantly in the Australasian region, though species are also found in other parts of the world such as Asia and South America. Most members of the family are epiphytic (growing on trees) or epilithic (growing on rocks). The composition of the family has varied significantly over the years, and as currently circumscribed the various Lembophyllaceae lack any reliable shared morphological characters and are united on the basis of molecular data (Olsson et al. 2009). Lembophyllaceae usually have concave leaves, loosely appressed to terete shoots, that lack a clearly differentiated leaf margin (Olsson et al. 2009).

In the broader context, Lembophyllaceae are placed among the pleurocarpous mosses of the order Hypnales (‘pleurocarpous’ means that the reproductive structures are produced on small side branches rather than at the top of the main stalk), probably as the sister group to the bulk of the Neckeraceae (Olsson et al. 2009). Merget & Wolf (2010) reported finding Lembophyllaceae as polyphyletic but a closer look at the supplementary figures shows that only two of their four clades (‘Lembophyllaceae III’ and ‘IV’) actually correspond to the current concept of the family established by Quandt et al. (2009); the remainder represent genera removed to other families. More to the point, Merget & Wolf were using that most wretched of molecular methods, neighbour joining, and despite their use of a large number of source taxa it is difficult to accord their results much significance.

One genus of Lembophyllaceae, Weymouthia, contains two species found in Australia, New Zealand and South America. Such disjunct distributions have been the subject of much debate in moss biogeography. Some authors attribute them to a Gondwanan ancestry, indicating that the modern species must have arisen prior to the separation of these landmasses through continental drift. Others would see them as the result of post-separation dispersal. Supporters of the former explanation point to the supposed inability of moss spores to survive extended environmental exposure, and the correlation of numbers of shared species with age of geological separation rather than current distance (Blöcher & Frahm 2002; e.g. South America shares more species with New Zealand than southern Africa, despite being closer geographically to the former). Supporters of the latter point to the low levels of morphological and molecular differentiation between individuals from disjunct populations.

Systematics of Lembophyllaceae

Characters (from Quandt et al. 2000): Leaves ovate-oblong, concave; rhombic apical cells; prorate, usually linear mid-laminal cells; differentiated basal and alar cells; short, mostly double costae. Exserted capsules on smooth setae; hypnoid peristome; calyptrae narrow, cucullate, mostly smooth.

<==Lembophyllaceae
    |--Fallaciella Crum 1991SK02
    |    `--F. gracilis (Hooker & Wilson) Crum 1991SK02 (see below for synonymy)
    |--Weymouthia Broth. 1906SK02
    |    |--W. cochlearifolia (Schwägr.) Dixon 1927 (see below for synonymy)SK02
    |    `--W. mollis (Hedw.) Broth. 1906 (see below for synonymy)SK02
    |--PorotrichodendronJ87
    |    |--P. bertrandiiJ87
    |    `--P. superbumJ87
    |--Lembophyllum Lindb. 1872SK02
    |    `--L. divulsum (Hooker & Wilson) Lindb. 1896 (see below for synonymy)SK02
    |         |--L. d. var. divulsumSK02
    |         `--L. d. var. clandestinum (Hooker & Wilson) Wijk & Margad. 1961SK02
    `--Camptochaete Reichardt 1870SK02
         |--C. angustata (Mitt.) Reichardt 1870 [=Isothecium angustatum]SK02
         |--C. arbuscula (Smith) Reichardt 1870 (see below for synonymy)SK02
         |--C. curvata Tangney 1997SK02
         |--C. deflexa (Wilson) Jaeger 1877 (see below for synonymy)SK02
         |--C. excavata (Taylor) Jaeger 1877 (see below for synonymy)SK02
         |--C. leichhardtii (Hampe) Broth. 1907 (see below for synonymy)SK02
         |--C. porotrichoidesBW15
         |--C. (Eucamptochaete) prolongata Brotherus & Watts 1915BW15
         `--C. pulvinata (Hooker & Wilson) Jaeger 1877 [incl. C. beckettii]SK02

Camptochaete arbuscula (Smith) Reichardt 1870 [=Hypnum arbuscula, Isothecium arbuscula, Porotrichum arbusculum, Thamnium arbuscula, Stereodon arbuscula; incl. Thamniella bateana (n. n.), Camptochaete longicaudata (n. n.)]^SK02

Camptochaete deflexa (Wilson) Jaeger 1877 [=Hypnum deflexum, Ca. arbuscula var. deflexa (Wilson) Dixon 1927, Stereodon deflexus, Thamnium deflexum; incl. Camptochaete brisbanica (nom. inv.), Thamniella brisbanica (n. n.), Thamniella campylochaete (nom. inv.), Hypnum chlorocladium, Coelidium chlorocladum, Lembophyllum chlorocladon, Thamnium curvatocomosum (n. n.), Camptochaete deflexula (n. n.), Thamnium deflexulum (nom. inv.), Isothecium fruticosum, Camptochaete fruticosa (Bastow) Paris 1900, Porotrichum fruticosum, Ca. nanodendra (nom. inv.), Thamnium deflexum var. perpumilum, Hypnum ramulosum (n. n.), Isothecium ramulosum, Camptochaete ramulosa (Mitt.) Jaeger 1877, Porotrichum ramulosum, Thamnium ramulosum, Camptochaete squamulosa (nom. inv.), Ca. squamulosula (nom. inv.)]^SK02

Camptochaete excavata (Taylor) Jaeger 1877 [=Hypnum excavatum, Thamnium excavatum; incl. H. assimile (n. n.), H. orbiculare Hampe 1880 (n. n.), Porotrichum orbiculare Mitt. 1882 (n. n.), Thamnium perpusillum (n. n.), Hypnum schlosseri, Brachythecium schlosseri, Camptochaete schlosseri Sendtn.) Broth. ex Paris 1909, Porotrichum schlosseri, Rigodium schlosseri, Thamniella schlosseri (n. n.), Hypnum vagum, Astrotrichum vagum (nom. inv.), Camptochaete vaga (Hornsch. ex Müll.Hal.) Broth. 1907, Coelidium vagum, Lembophyllum vagum, Porotrichum vagum, Rigodium vagum, Thamniella vaga, Thamnium vagum]^SK02

Camptochaete leichhardtii (Hampe) Broth. 1907 [=Hypnum leichhardtii, Ptychomnion leichhardtii, Thamniella leichhardtii, Thamnium leichhardtii]^SK02

Fallaciella gracilis (Hooker & Wilson) Crum 1991^SK02 [=Isothecium gracile^SK02, Camptochaete gracilis (Hooker & Wilson) Paris 1894^SK02, Microthamnium gracile^SK02, Porotrichum gracile^SK02, Stereodon gracilis^SK02, Thamnium gracile^SK02; incl. Lembophyllum brisbanicum (n. n.)^SK02, Porotrichum brisbanicum Müll.Hal. ex Bailey 1883 (n. n.)^SK02, Camptochaete tasmanica^SK02]

Lembophyllum divulsum (Hooker & Wilson) Lindb. 1896 [=Hypnum divulsum, Coelidium divulsum, Lembophyllum clandestinum var. divulsum (nom. illeg.), Porotrichum divulsum, Thamniella divulsa]^SK02

Weymouthia cochlearifolia (Schwägr.) Dixon 1927 [=Hypnum cochlearifolium, Coelidium cochlearifolium, Isothecium cochlearifolium, Lembophyllum cochlearifolium, Porotrichum cochlearifolium, Stereodon cochlearifolius; incl. Neckera billardieri, Meteorium billardieri, Pilotrichella billardieri, Weymouthia cochlearifolia var. billardieri (Hampe) Dixon 1927, Hypnum cochlearifolium var. minus Hampe 1853 (n. n.) nec H. collatum var. minus nec H. cupressiforme var. minus nec H. polygamum var. minus, Coelidium cochlearifolium var. minus Paris 1894 (n. n.), Lembophyllum cochlearifolium var. minus Paris 1896]^SK02

Weymouthia mollis (Hedw.) Broth. 1906 [=Leskea mollis, Isothecium molle, Meteorium molle, Neckera mollis, Pilotrichella mollis, Stereodon mollis; incl. Meteorium molle var. majus, Pilotrichella mollis var. major (Wilson) Jaeger 1877, P. weymouthii]^SK02

*Type species of generic name indicated

References

Blöcher, R., & J.-P. Frahm. 2002. A comparison of the moss floras of Chile and New Zealand. Tropical Bryology 21: 81–92.

[BW15] Brotherus, V. F., & W. W. Watts. 1915. The mosses of the New Hebrides [part 1]. Journal and Proceedings of the Royal Society of New South Wales 49 (1): 127–144.

[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum—Biological Sciences 32 (1): 1–136.

Merget, B., & M. Wolf. 2010. A molecular phylogeny of Hypnales (Bryophyta) inferred from ITS2 sequence-structure data. BMC Research Notes 3: 320.

Olsson, S., V. Buchbender, J. Enroth, S. Huttunen, L. Hedenäs & D. Quandt. 2009. Evolution of the Neckeraceae (Bryophyta): resolving the backbone phylogeny. Systematics and Biodiversity 7 (4): 419–432.

Quandt, D., S. Huttunen, R. Tangney & M. Stech. 2009. Back to the future? Molecules take us back to the 1925 classification of the Lembophyllaceae (Bryopsida). Systematic Botany 34 (3): 443–454.

Quandt, D., R. S. Tangney, J.-P. Frahm & W. Frey. 2000. A molecular contribution for understanding the Lembophyllaceae (Bryopsida) based on noncoding chloroplast regions (cpDNA) and ITS2 (nrDNA) sequence data. Journal of the Hattori Botanical Laboratory 89: 71–92.

[SK02] Streimann, H., & N. Klazenga. 2002. Catalogue of Australian Mosses. Flora of Australia Supplementary Series 17. Australian Biological Resources Study: Canberra.