Barbula consanguinea (Thw. & Mitt.) Jaeg. sensu Eddy is considered as a pan-tropical species. This taxon is reported new for several central African countries, the Arabian peninsula, and Middle America. This variable species is shortly discussed and compared with (most) related taxa. Hymenostylium crispulum Broth. & Par. and Barbula obscura Sull. (= Barbula wrightii Sauerb.) are considered as synonyms of Barbula consanguinea

Sollman, Philip

English

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17TROPICAL BRYOLOGY 19 (2000)Studies on Barbula consanguineaTropical Bryology 19: 17-23, 2000Studies on Barbula consanguinea (Thw. & Mitt.) Jaeg. sensu Eddy,a pan-tropical speciesPhilip SollmanVon Weberstraat 32, 6904 KD Zevenaar, The NetherlandsAbstract. Barbula consanguinea (Thw. & Mitt.) Jaeg. sensu Eddy is considered as a pan-tropicalspecies. This taxon is reported new for several central  African countries, the Arabian peninsula, andMiddle America. This variable species is shortly discussed and compared with (most) related taxa.Hymenostylium crispulum Broth. & Par. and Barbula obscura Sull. (= Barbula wrightii Sauerb.) areconsidered as synonyms of Barbula consanguinea.IntroductionOver the last few years, the author has studiedmany herbarium collections, mainly held in L,which belong to Barbula consanguinea, mostlycollected in tropical Asia.Eddy (1990: 178) realized that Barbulaconsanguinea is a (very) variable species.Furthermore, the species is rather commonlydistributed, at least in tropical Asia.Type material of this taxon was also studied, viz.the isotypes present in BM, GRO, H.  In severalcases, the isotype material consisted of mixturesof two (related) taxa, viz. Barbula javanica Dozy& Molk. and Barbula consanguinea sensu Eddy(Eddy 1990).Furthermore, the isotype material in the Brotherusherbarium (H) is very sparse. This collection wasannotated by B.C. Tan in 1983 as Barbula indica(Hook.) Spreng. However, in my opinion, thismaterial is better referred to a smaller (younger)state of Barbula consanguinea (sensu Eddy).In addition, the holotype of Barbulaconsanguinea was studied (NY!). I agree withSaito (1975: 495), that the plants belong toBarbula javanica Dozy & Molk. No admixtureis present in the studied holotype material.However, the concept presented in the Asianfloras of Bartram (1939), Chen (1941), Fleischer(1904), Gangulee (1972) is consistent anddifferent. This is the reason why I follow this line,also present in Eddy’s flora, so sensu Eddy.For the moment, Barbula consanguinea is theoldest name for several taxa which are closelyrelated, in my opinion idential.Material and methodIn total, about 200 collections were studied, filedand often annotated. All the material present inherbarium L was examined.Furthermore, most plants were studied severaltimes.In addition, selected material from the herbaria18TROPICAL BRYOLOGY 19 (2000)SollmanBM, CANB, H-BR., MELU-Stone, NSW, NYwas inspected.Various type collections, belonging to this -Hydrogonium-group, were also examined.All the material was studied wet under themicroscope. Field survey on this species, weredone in 1993 in Australia (Queensland) and thePhilippines (Luzon).Description, discussion, figuresThis species is well described, discussed andillustrated especially by Eddy (1990: 178-180).Furthermore, Barbula consanguinea is welldescribed and figured by Fleischer (1904: 348-350), although no gemmae are mentioned.Hilpert (1933: 625, 627) discusses Barbulaconsanguinea shortly.The species is not illustrated here.Bartram (1939: 119) gave a good, but rather shortdescription.The figure (plate 9, fig. 142) is very concise.Chen (1941: 240, Abb. 45, 1-3) gave only a shortdiscussion and a plate. However, in the text belowseveral -Hydrogonium- taxa, Chen (l.c.)compared several species with each other andwith Barbula consanguinea.Gangulee (1972: 729) gave a good descriptionand figure.Saito (1975: 493) gave mainly a good description,illustration and discussion as Barbula subcomosaBroth. However, in his description (p. 493) thephrases “laminal cells ... rarely smooth ...” and“costa ... ending below apex ...” are notappropriate for this taxon. In Saito (1972: 11)the observed variation on the gemmae isespecially discussed and illustrated (also asBarbula subcomosa). Furthermore, in myopinion, Barbula subcomosa Broth. is identicalwith Barbula consanguinea sensu Eddy.In Noguchi (1988: 299-301) Barbula subcomosaBroth. is described as laminal cells “...mammillose ...”(idem in the key and also fig. 125,A). However, this is not correct. The laminal cellsin this taxon are clearly pluripapillose.     Norris and Koponen (1989: 118-119) listedBarbula consanguinea as a synonym belowBarbula javanica. They followed Saito (1975:495), without any further comment. In theirdiscussion (l.c. p. 119) below Barbula javanica,they suggested that “... Barbula javanica isquestionably distinct from Barbula subcomosa...”.However, in my opinion, both taxa are clearlydistinct. In Barbula javanica the laminal cells areclearly mammillose, the cells pellucid. In Barbulasubcomosa, on the contrary, the laminal cells areclearly pluripapillose and obscure, especially inthe upper parts of the (upper) leaves, among otherdifferences between these two taxa.The description, key characters and also theillustration (fig. 9, 1-o) in Norris and Koponen(1989) clearly refer to Barbula javanica Dozy &Molk.IllustrationsAkiyama (1996), p. 159, fig. 1; Bartram (1939),plate 9, fig. 142; Chen (1941) p. 240, fig. 45: 1-3, as Hydrogonium consanguineum; Gao (1996),p. 208, fig. 71: 1-3, as Hydrogoniumconsanguineum; Eddy (1990), p. 179, fig. 278;Fleischer (1904), p. 349, fig. 63; Gangulee(1972), p. 733, fig. 348, as Hydrogoniumconsanguineum.VariationMost variation was seen in:1. general tingeThe plants are commonly tan coloured, morerarely (dull) green in the upper part.2. heightCommonly about 1.5 cm. Ranging from c. (0.5)1 to 3 cm, rarely larger (up to c. 5 cm).3. general leaf outlineCommonly lingulate to triangular. The betterdeveloped leaves ofter tapering from near baseto apex; in younger leaves not so pronounced.4. leaf lengthCommonly about 2 mm long. Ranging fromrather small to longer c. 1.5 - 2.2 mm. Thevariation here is sometimes considerable even inone tuft.5. apical region of the leavesCommonly circa cuspidate. Ranging frombroadly rounded (mucronate) to small acute. Thevariation can be present in one tuft, more rarelyeven on one plant.6. narrowly revolute leaf margins just above leafbaseCommonly narrowly recurved just aboveleafbase, at least in some leaves; more rarely19TROPICAL BRYOLOGY 19 (2000)Studies on Barbula consanguineanarrow recurved to about halfway up (in someleaves). Rather rarely most leaves plane or nearlyso.7. density of the laminal papillae in the upperpart of the (upper) leaves.The laminal cells in this region are clearly denselypluripapillose, obscuring the lumen. However,there is some variation present - especiallybetween collections- in the density of the laminalpapillae.8. excurrent part of the costaCommonly shortly excurrent: (1) 2-4 smoothcells. Sometimes hardly excurrent on severalleaves and/or broken, especially on older leaves.9. extreme leaf apex (entire- indented- somewhatdenticulate)Commonly entire to somewhat indented or(slightly) denticulate in this region. However, thiscondition is often variably expressed incollections examined.10. back of the costaCommonly strongly papillose and also partlydouble papillose, from apex to near base. Morerarely, the papillae are not so pronounced (onsome leaves).11. axillary gemmaeThey are constantly present, although sometimesscarce. Commonly stalked ovoid, ellipsoid,clavate, or fusiform, multicellular, greenish to(dark) brownish with age. More rarely alsolocally bulging/knobby, to forked/branched.Part of the observed variation is discussed andillustrated by Saito (1972:11), as Barbulasubcomosa Broth.Interesting collection studied:Australia, Queensland, (W. of Cairns), Mungana,on earth, near mine, 19 Aug. 1979, I.G. Stone15.933, det. Ph. Sollman, Jan. 1999, hb. MELU.This number shows a remarkable variation ingemmae shapes.Barbula  indica Barbula consanguineageneral height: to about plants commonly larger, to about1 cm; small plants 3 cm; seldom somewhat highergeneral tinge: commonly commonly tan coloureddull (darker) greenleaf margins parallel leaf margins commonly taperingfrom near base to apexleaf margins often narrowly leaf margins often only slightlyrecurved from near base to narrowly recurved below, at leasthalfway, or more in some (larger) leaves; sometimesalmost planeleaf apex broadly rounded leaf apex commonly more acute(acuminate-cuspidate), in some younger states often more broadlyroundedexcurrent point of the costa excurrent point of the costavery short, mostly 1 cell often, more pronounced: (1) 2-4cellsperistome teeth about straight peristome teeth clearly twistedTab. 1: Differences between Barbula indica and B. consanguinea.20TROPICAL BRYOLOGY 19 (2000)Sollman12. fragility of the leavesMore or less fragile plants occur rather regularly,especially the older parts. Sometimes thiscondition is rather pronounced, with almost onlythe costa intact.Plants encrusted with lime occur also now andthen.ComparisonSome, small non-fruiting states of Barbulaconsanguinea resemble Barbula indica (Hook.)Spreng., sometimes rather closely. Using tab. 1scheme,one can separate difficult collections.Furthermore there are habitat differences betweenthe two taxa. Barbula indica very often grows inbuilt-up areas, on stone, e.g., boulders, brickwork,masonry etc. often with Hyophila involuta(Hook.) Jaeg. (nearby), regularly rather dry.Barbula consanguinea, on the contrary, growsin more natural habitats (e.g., boulders, rocks,etc. along water), often (much) wetter thanBarbula  indica.Some states of Barbula consanguinea resembleBarbula unguiculata Hedw., sometimes ratherclosely. However, the latter lacks axillarygemmae, which are constantly present in Barbulaconsanguinea, although sometimes scarce.Furthermore, there are clearly differences in thedensity of the papillae on the back of the costa.In Barbula unguiculata they are (very) low,mostly present in the apex region (only). On thecontrary, in Barbula consanguinea they arecommonly far more pronounced, from apex tonear base.     In addition, the geographical ranges of bothtaxa are different. Barbula unguiculata is mainlypresent in the temperate region. Barbulaconsanguinea is mainly a pantropical species.However, in some regions their ranges can comeclose or overlap partly, e.g., parts of Australia,China, Japan, USA. Furthermore, Barbulaunguiculata is (sometimes) introduced incountries with soil.In Australia, Barbula consanguinea has beenconfused sometimes with Barbula subcalycinaC. Muell.  For a discussion on this species, seeStone (1990: 265-266).In non-fruiting collections of Barbulasubcalycina, the costa is (far) stronger, thelaminal papillae (far) more pronounced (Tortellatype), tomentum is present and the details of theleaf base are different. Furthermore, gemmae arenot known in Barbula subcalycina.In tropical America it is possible that especiallyBarbula orizabensis C. Muell. is mistaken forBarbula consanguinea.Tab 2 will separate the two taxa.Barbula seramensis H. Akiyama is also nearBarbula orizabensis Barbula consanguineagemmae commonly globose, gemmae commonly stalked ovoidsmall to club-shaped, often alsofusiform, largerexcurrent part of the costa: commonly only weakly excurrentclearly shortly excurrent (1) 2-4 cells, smoothleaf margins recurved from near leaf margins only narrowlybase to apex recurved in lower part of theleaves, rarely to about halfwayTab. 2: Differentiationof Barbula orizabensis and B. consanguinea.21TROPICAL BRYOLOGY 19 (2000)Studies on Barbula consanguineaBarbula consanguinea.For a discussion and illustration, see especiallyAkiyama (1996).RecognitionI recognize Barbula consanguinea as follows:1. the leaf outline commonly tapering from abroader base to a smaller (often cuspidate) apex;2. the laminal cells in the upper part of the (upper)leaves clearly densely pluripapillose, obscuringthe lumina;3. leaf margins often narrowly recurved below,just above leaf base, at least in some (larger)leaves;4. gemmae are constantly present, althoughsometimes scarce;5. back of the costa commonly papillose to partlydouble papillose, from apex to near base;6. costa commonly shortly excurrent, about (1)2-4 cells, at least in some (larger) leaves;7. peristome teeth twisted.HabitatThis species grows especially in areas with -atleast some- lime and water present, or nearby.Along water, (seasonal) rivers,  rivulets, creeks,mudpools, gullies, springs, near waterfalls(sprayzone), cascades, cataracts. On stonysubstrata with -at least- some soil covering,permanently, or periodically moist to wet, suchas: (lime) stone, cliffs, boulders, travertine,(dripping) rocks, concrete, sandstone, (concreted)walls, gorge walls. Furthermore, also terrestrialon soil, sand, clay, mud,  gravel, (compacted)banks, along tracks, roadsides, along rice-fields,soil walls, waterchannels.This species grows sometimes (partly)submerged and is then often encrusted with lime;rarely also tufa-forming. Very rarely on tree bases-with some soil cover- e.g., coconut.In tropical Asia commonly growing in partialshaded situations to more open, exposed, morerarely in heavy shade.AltitudeMost collections studied came from sealevel toabout 1000 m., more seldom (somewhat) higher.New reportsBarbula consanguinea is reported new for:Central AfricaBurundi, route de Bujumbura à Rumonge, prèsde Kitanza, mousse incrustante dans un tuf aubord de la route, sterile, with gemmae, alt. c. 800m., 16 Sept. 1974, J.L. de Sloover 19.200, det.Ph. Sollman, March 1997, hb. L, de Sloover.Notes.1. In herbarium present as Hydrogonium spec.Cape Verde IslandsBrava, Ribeira da Faja da Agua, an feuchter, sehrsandiger Böschung, W. exponiert, sterile, withgemmae, alt. 35-50 m., 28 Jan. 1995, A. Lindlar68 + 70, det. Ph. Sollman, Jan. 1997, hb. L, DUIS.Notes.1. These collections were also listed in Frahm etal. (1996), as cf identifications.GuineaGuinea gall., cercle de Kourroussa, pr. pagumBanka, ad rupes fontium, sterile, with gemmae,Jun. 1903, H. Pobeguin s.n., ex hb. E.G. Paris, L910.115.172.Notes.1. This collection is isotype material ofHymenostylium crispulum Broth. & Par. Thematerial agrees well with the description in theprotologue (Revue Bryol. 31: 42. 1904). SomeHyophila involuta as admixture is present. Noother admixture was found.2. This collection agrees well with Barbulaconsanguinea (Thw. & Mitt.) Jaeg. sensu Eddy.NigeriaNiger Prov., Bonu Abuja Distr., waterfalls on theriver Gurara, on hard steep rocks, in dry channelsthat carry water in times of flood, sterile,s. alt.,19 April 1958, E.W. Jones 994-A, ex hb. C.C.Townsend, det. Ph. Sollman, Dec. 1996; idemJan. 1997, hb. L.Notes.1. Material separated from Hyophila involuta. A-number given by the author.Arabian peninsulaOman, on rocks, in flowing water, Wadi Hinna,below the Baobab car park, sterile, with gemmae,alt. c. 200 m., lat. 17° 03’N, 54° 36’E, 24 Sept.1992, H.D.V. Prendergast B 33, B 38, det. Ph.Sollman, 1997; hb. Townsend, L.22TROPICAL BRYOLOGY 19 (2000)SollmanNotes.1. Part of the material with fragile/broken leaves.Oman, on rocks, shaded, Wadi Tobruk, sterile,with gemmae, alt. c. 100 m., lat. 17° 05' 47"N,054° 19' 37" E, 25 Sept. 1992, H.D.V.Prendergast B 40, det. Ph. Sollman, 1997, hb.Townsend, L.Tropical Middle AmericaCuba, dry banks, Wright 31, det. Ph. Sollman,1993, hb. L 909. 335.93.Notes.1. This is an isotype collection of Barbulaobscura Sull. (= B. wrightii Sauerb.). See alsoZander, 1979: 186. Zander (l.c.) treats this taxonas Barbula indica (Hook.) Spreng. However, thiscollection agrees well -in my opinion- withBarbula consanguinea material studied fromtropical Asia.2. Gemmae are present.USAFlorida, Walton County, Lake Wood Park,highest point in Florida, c. 0.7 m. S. of Alabama- Florida State line, along Walton Co. road 1087,on calcareous soil along road, alt. 345 ft, fruiting,with gemmae, 30 Nov. 1988, B.H. Allen 7541,det. Ph. Sollman, 1999, hb. L 988.051.332, MO.Notes.1. Det. B. Allen, 1989 as Barbula unguiculataHedw.Florida, Jackson County, Florida Caverns StatePark, three miles north of Marianna, on groundin a path in woods along Chipola River, sterile,with gemmae, s. alt., 4 Jan. 1949, F. Drouet, C.S.Nielsen c.s. (s.n.?), det. Ph. Sollman, 1999, hb. L952.83. 197.Notes.1. Det. M. Feigley, 1950 (no. 10.348) as Barbulaunguiculata Hedw.Hawaii, Oahu, Waianae Mts., summit KoalaMetrosideros -Cheirodendron moss forest,partially submerged in intermittent roadsidepuddles, sterile, with gemmae, alt.c. 4000 ft, 22Feb. 1975, W.J. Hoe 3347.0, det. Ph. Sollman,1999, hb. L 976.198.657.Notes.1. Det. K. Saito as Barbula unguiculata Hedw.2. Bryophyta Hawaiica Exsic., Ed. by W.J. Hoe,Ser. I: Musci no. 22. Barbula unguiculata Hedw.AustraliaTropical parts of Australia, viz. Queensland,Northern Territory, W. Australia. About 60collections were studied. The material is mainlypresent in the herbaria BM, CANB, H-BR., L,NSW, MELU-Stone.LiteratureAkiyama, H. 1996. Taxonomic studies of mosses ofSeram and Ambon (Moluccas, East Malesia)collected by Indonesian - Japanese BotanicalExpeditions, IX Pottiaceae – Tropical Bryology12: 157-168.Bartram, E.B. 1939. Mosses of the Philippines -Philippine J. Science 68: 1-147.Bartram, E.B. 1949. Mosses of Guatemala. FieldianaBot. 25: I-V, 1-442.Chen, P.C. 1941. Studien über die ostasiatischen Artender Pottiaceae I, II - Hedwigia 80: 1-76; 141-322.Chuang, C.C. 1973. A Moss Flora of Taiwanexclusive of essentially pleurocarpous Families -J. Hattori Bot. Lab. no. 37: 419-509.Eddy, A. 1990. A handbook of Malesian Mosses, vol.2. London.Fleischer, M. 1904. Die Musci von Buitenzorg, ErsterBand. Leiden.Frahm, J.P. et al. 1996. Bryophytes from the CapeVerde Islands - Tropical Bryology 12: 123-153.Gangulee, H.C. 1972. Mosses of Eastern India andadjacent regions, Fasc. 3. Calcutta.Gao, C. (ed.) 1996. Flora Bryophytarum Sinicorum,vol. 2. Fissidentalis - Pottiales. Science Press,Beijing (in Chinese).Hilpert, F. 1933. Studien zur Systematik derTrichostomaceen - Beih. z. Bot. Centralbl. 50, Abt.II: 585-726.Miller, H.A., H.O. Whittier & B.A. Whittier, 1978 .Prodromus Florae Muscorum Polynesiae, with akey to genera. J. Cramer, Vaduz.Noguchi, A. 1988. Illustrated Moss Flora of Japan,Part 2,  Supplemented by Z. Iwatsuki. Nichinan.Norris, D.N. & T. Koponen, 1989. Bryophyte floraof the Huon Peninsula, Papua New Guinea XXVIIIPottiaceae (Musci) - Acta     bot. Fennica 137: 81-138.Saito, K. 1972. Notes on the Pottiaceae (2) - Journ.Jap. Bot. Vol. 47, no. 1: 10-19.Saito, K. 1975. A monograph of Japanese Pottiaceae(Musci) - J. Hattori bot. Lab. 39: 373-537.23TROPICAL BRYOLOGY 19 (2000)Studies on Barbula consanguineaStone, I.G. 1990. Nomenclatural changes and newmoss records in Australia: including a descriptionof the protonema of Calomnion - J. Bryol. 16: 261-273.Zander, R.H. 1979. Notes on Barbula andPseudocrossidium  (Bryopsida) in North Americaand an annotated key to the taxa - Phytologia vol.44, no. 4: 177-214.Zander, R.H. 1994 Pottiaceae in: A.J. Sharp et al.The Moss Flora of  Mexico, New York.24TROPICAL BRYOLOGY 19 (2000)Sollman

Studies on Barbula consanguinea (Thw. & Mitt.) Jaeg. sensu Eddy : a pan-tropical species

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