Classification
Class
 Nomenclature
Scientific Name:
Sematophyllaceae Broth., Nat. Pflanzenfam. [Engler & Prantl] 1(3), 706 (1905)
 Description

Plants slender to robust, forming dense mats or tufts, often glossy. Stems creeping or ascending, irregularly to densely and pinnately branched, without a central strand, usually ± red. Stem and branch leaves similar or occasionally differentiated (as in Wijkia), crowded in many rows but sometimes complanate, often homomallous or secund, mostly symmetric, variously shaped but often ± ovate and often acuminate (rarely broad or rounded) at apex, mostly variably toothed, mostly unbordered. Laminal cells linear to rhombic, smooth or papillose; alar cells nearly always clearly differentiated, often inflated, vesiculose at extreme angles, less often ± quadrate. Costae short and double or lacking. Paraphyllia lacking. Pseudoparaphyllia often present and foliose.

Autoicous or dioicous. Perichaetia lateral and with the leaves differentiated. Setae elongate, mostly smooth but sometimes papillose; capsules mostly inclined to pendulous and asymmetric, rarely erect and symmetric (in non-N.Z. taxa), ovoid- or oblong-cylindric, smooth, mostly pale brown; exothecial cells often collenchymatous or subcollenchymatous; stomata restricted to neck; annulus often not differentiated; operculum conic at base, short- or long-rostrate. Peristome mostly double and hypnoid, pale. Calyptra cucullate or rarely mitrate, smooth. Spores spherical.

 Taxonomy

The Sematophyllaceae are a large and predominantly tropical and subtropical family with limited diversity in temperate regions. Brotherus (1925) recognised 36 genera distributed worldwide and placed them in four subfamilies.

The family is taxonomically difficult in most parts of its range and even the modest number of taxa occurring regionally pose some practical identification difficulties.

The generic limits in the Sematophyllaceae remain highly controversial and fluid. They are likely to become more settled only after large-scale monographic study, including molecular analysis, of this family. Hedenäs & Buck. W.R. (1999) performed a cladistic analysis of the genera traditionally placed in the family using morphological characters and recognised 35 genera (their Table 7), three of which they placed in a subfamily Wijkioideae. They suggested that these three genera (Wijkia, Acanthorrhynchium, and Trismegistia) are the "genera of Sematophyllaceae that have the least specialised perichaetia and sporophytes, and … likely to be the most primitive ones in the family." This result largely mirrors the conclusions of Tan & Jia (1998) in their cladistic study of Chinese members of the family. Ten years later, Goffinet et al. (2009) proposed a radically different classification, recognising (within the Hypnales) a Sematophyllaceae of 28 genera while simultaneously placing several genera traditionally assigned here in a newly described family, the Pylaisiadelphaceae. Among the genera they placed in the latter family were Taxithelium, Trismegistia, and Wijkia.

Even the eponymous genus Sematophyllum is poorly circumscribed, with Buck (1998, p. 368) describing it as "scarcely more than the dregs of the Sematophyllaceae… characterized by the lack of various specialised features."

The Australian flora, with it tropical components, is far richer than the N.Z. flora for this family. Ramsay et al. (2002; 2004) provided an Australian treatment recognising 14 genera that helps to place our more limited flora in context. Their discussion of large-scale systematic modifications proposed to the family prior to 2002 also provided some clarity to a confusing set of proposals. Because of the lack of consensus concerning the limits of the Sematophyllaceae, and generic boundaries within it, it is expedient to adopt the generic assignments proposed by Ramsay et al. (2002; 2004). For their regional revision Ramsay et al. employed a largely Brotherean classification, albeit with several exceptions, including notably: (1) they did not recognise the four subfamilies employed by Brotherus (1925) and (2) they excluded from the family the large and well-known genus Taxithelium (which does not occur in N.Z.). Tan et al. (1996) presented a useful key to the Australian genera.

Four genera, nine species, and one non-typical variety are accepted as part of the N.Z. flora. While generic placements largely follow Ramsay et al. (2002; 2004), not all of their species-level taxonomy is accepted here. For reasons alluded to above, no attempt is made to present a key to the N.Z. genera.

 Key
1Stem and branch leaves differentiated in size and shape; stem leaves abruptly tapered to a piliferous and serrulate apex; laminal cells either pluripapillose or smooth; operculum high-conic, lacking a rostrum 2
1'Stem and branch leaves not or scarcely differentiated; stem leaves lacking a piliferous and serrulate apex; laminal cells smooth; operculum with a conic base and a long, slender rostrum 3
2Stems irregularly or subpinnately (never bipinnately) branched; branches simple and erect or curving upwards from the substrate, weakly cuspidate and with numerous flagelliform and microphyllous branchlets near tips; branch leaves entire or crenulate near apex; mostly 0.7–0.8(–1.1) mm long Wijkia extenuata var. caudata
2'Stems usually bipinnately branched; branches simple or subpinnately branched, often tapered to a slender and cuspidate tip, lacking flagelliform branchlets; branch leaves sharply serrulate above by projecting cell ends (usually more serrulate than stem leaves), 1.1–1.4 mm long Wijkia extenuata var. extenuata
3Dioicous; plants nearly always coastal, gold-brown or copper in colour; leaves strongly homomallous and pointing away from the substrate; alar group large, each extending laterally ¼ to nearly ⅓ the total width of leaf and extending up margin for 6–10 cells, composed of firm-walled, subquadrate, and opaque cells or with c. 3–4 cells in the extreme basal angle moderately enlarged and surrounded by firm-walled opaque cells; capsules suberect Sematophyllum homomallum
3'Autoicous or sexuality unknown; plants not restricted to coastal situations, various in colour; leaves variously oriented (strongly homomallous and pointing away from substrate only in S. subhumile var. contiguum); alar group smaller, with the cells at extreme corners thin- or firm-walled, enlarged, not opaque; capsules horizontal to pendulous, rarely inclined, or unknown4
4Leaves very long and fine, mostly 3.5–4.5 mm, strongly falcate-secund; sexuality unknown (neither sex organs nor capsules known); rare and known from Southland L.D. and Stewart I. Sematophyllum fiordensis
4'Leaves much shorter, ≤2.7 mm, orientation various (sometimes falcate-secund); autoicous and often with sporophytes; widely distributed 5
5Leaves evenly tapered to a slender and acuminate apex, mostly 0.2–0.4 mm wide (under cover slip) 6
5'Leaves either unevenly tapered or broadly acute at apex, usually >0.4 mm wide (under cover slip)8
6Vegetative leaves sharply denticulate above, erect-appressed or weakly homomallous; setae mammillate; exothecial cells subcollenchymatous and in distinct longitudinal ranks; known only from southernmost N.Z. (Moggy, Solander, The Snares, and Auckland Is) Sematophyllum kirkii
6'Vegetative leaves entire or weakly denticulate above, either falcate-secund (in Rhaphidorrhynchium amoenum) or strongly homomallous (in Sematophyllum subhumile var. contiguum); setae smooth; exothecial cells clearly collenchymatous, not in distinct longitudinal ranks; widespread in N.Z. 7
7Plants caespitose; leaves homomallous, pointing away from the substrate, entire; setae 7–9 mmSematophyllum subhumile var. contiguum
7'Plants forming interwoven mats, not caespitose; leaves strongly falcate-secund to nearly circinate, entire or finely denticulate; setae 8–20 mm Rhaphidorrhynchium amoenum
8Plants usually bright green and very lustrous; branches flattened and cuspidate apically; leaves elliptic and evenly tapered to a broadly acute apex, ± spreading or homomallous at branch tips, with numerous short apical cells; setae 6–11 mm, mammillate-papillose near apex (occasionally obscurely so); restricted to stream margins Sematophyllum jolliffii
8'Plants usually golden to yellow-brown, moderately lustrous or dull; branches neither flattened nor cuspidate apically; leaves ± ovate-lanceolate and ± abruptly tapered to an acute or acuminate apex, secund to ± falcate at branch tips, with only a few apical cells shortened; setae nearly always >10 mm, either smooth or mammillate; habitat various (only S. uncinatum restricted to stream margins) 9
9Plants relatively robust, hygrophilous, mostly saxicolous or on thin soil over rock (only occasionally on wood) and restricted to stream beds; leaves broadly ovate-lanceolate, 0.45–0.85 mm wide (under cover slip) Sematophyllum uncinatum
9'Plants fine, not hygrophilous, mostly lignicolous or epiphytic; leaves narrowly ovate-lanceolate, c. 0.20–0.35 mm wide (under cover slip) 10
10Setae mammillate; capsule neck irregularly protuberant/mammillate, with stomata situated at the apex of irregular protuberances; exothecial cells subcollenchymatous, arranged in distinct longitudinal ranks (visible under hand-lens); leaves mostly 0.30–0.35 mm wide (under cover slip) Warburgiella leucocyta
10'Setae smooth; capsule neck weakly differentiated, lacking irregular protuberances; exothecial cells clearly collenchymatous, not in longitudinal ranks; leaves narrower, mostly 0.20–0.30 mm wide (under cover slip) Rhaphidorrhynchium amoenum
 Biostatus
Indigenous (Non-endemic)
Number of species in New Zealand within Sematophyllaceae Broth.
CategoryNumber
Indigenous (Endemic)2
Indigenous (Non-endemic)7
Total9
 Excluded Taxa

Hypnum cyparioides Brid. According to Dixon, the type of this obscure name was collected by de Labillardière in "Nova Hollandiae". Both Dixon (1929, p. 308) and Ramsay et al. (2004) placed Hypnum cyparioides Brid. in the synonymy of Rhaphidorrhynchium amoenum (Hedw.) M.Fleisch. (or its nomenclatural equivalent). Mitten (1859) applied the name H. cyparioides to a Tasmanian collection by Archer. Dixon discussed the confusion between H. cyparioides Brid. and H. amoenum Hedw. at some length but could locate types of neither name. Hypnum cyparioides Brid. (or combinations based on it) have received little or no application to N.Z. collections and this name is not considered further.

Hypnum leptorhynchum Brid. This name seems to have been first discussed in relation to H. leucocytus Müll.Hal. and H. cerviculatum Hook.f. & Wilson by Dixon (1929, p. 309). The name H. leptorhynchum Brid. is used by Wilson & Hooker in both Flora Antarctica (1845, p. 141) and in Flora Novae-Zelandiae (Wilson 1854, p. 112) for material from N.Z. Confusingly, Wilson (1854, p. 113) described the setae in such material as smooth ("laevi"). Many of the older collections in BM named as H. leptorhynchum Brid. are referable to S. amoenum. According to Tropicos (accessed 2 Nov. 2015) H. leptorhynchum Brid. is an invalid name and it is discussed no further here.

Rhaphidostegium dallii Broth. & Geh. This species was described from a single collection of unlocalised N.Z. provenance. It is discussed by Dixon (1929, p. 309) who saw only vegetative material. He tentatively accepted it while stating that "vegetatively it agrees exactly with R. leucocytus." Sainsbury (1955, p. 466) listed it as a doubtful species. Ramsay et al. (2004, p. 58) examined type material and considered it to be a synonym of Warburgiella macrospora, which is considered here to be inseparable from W. leucocytus. It is not considered further here.

Sematophyllum subcylindricum sensu Sainsbury and sensu Allison. Some N.Z. specimens named (mostly by K.W. Allison) as S. subcylindricum (Broth.) Sainsbury in N.Z. herbaria are perplexing due to their poor quality and for having ± "subcollenchymatous" exothecial cells and smooth setae (e.g., G.O.K. Sainsbury 4740 from Hopuruahine Hill near Lake Waikaremoana, Gisborne L.D. (CHR 570159) and K.W. Allison 3207 from Pukerimu Bush, near Taupō, S Auckland L.D. (CHR 570161). Most material so-named is aberrant and poor Rhaphidorrhynchium amoenum. Sematophyllum subcylindricum is considered here to be worthy neither of taxonomic recognition nor of further consideration. The relevant collections remain filed under S. subcylindricum in CHR.

 Bibliography
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Dixon, H.N. 1929: Studies in the bryology of New Zealand, with special reference to the herbarium of Robert Brown. Part VI. Bulletin, New Zealand Institute 3(6): 299–372.
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Wilson, W.; Hooker, J.D. 1845 ("1847"): Musci. In: Hooker, J.D. The Botany of the Antarctic Voyage of H.M. Discovery Ships Erebus and Terror in the Years 1839–1843, under the command of Captain Sir James Clark Ross. I. Flora Antarctica. Part I. Botany of Lord Auckland’s Group and Campbell’s Island. Reeve, Brothers, London. 117–143.