The Genus Pseudocalliergon in Northern Europe

Oikos Editorial Office

The Genus Pseudocalliergon in Northern EuropeAuthor(s): Lars HedenäsSource: Lindbergia, Vol. 16, No. 3 (1990), pp. 80-99Published by: Oikos Editorial OfficeStable URL: http://www.jstor.org/stable/20149763 .

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LINDBERGIA 16: 80-99. Copenhagen 1990

The genus Pseudocalliergon in northern Europe

Lars Heden?s

9

Heden?s, L. 1992. The genus Pseudocalliergon in northern Europe. -

Lindbergia 16: 80-99.

The genus Pseudocalliergon (Limpr.) Loeske is revised for northern Europe. The

genus is interpreted as consisting of P. angustifolium Heden?s, sp. nov., P. lycopo dioides (Brid.) Heden?s, comb, nov., P. brevifolium (Lindb.) Fleden?s, comb, nov., P. turgescens (T. Jens.) Loeske and P. trifarium (Web. & Mohr) Loeske. The genus is defined by, i.a., the typical colour and gloss of the species, the structure of the alar cell group, the structure and colour of the axillary hairs and the ecology of the

species. All the species grow in lime-rich and, except for P. trifarium, small and often shallow and periodically dry wetland habitats (sometimes the habitats are flushed

with ?lime-rich water). In northern Europe, P. angustifolium grows in the Scandina vian mountain range and in the more eastern areas also in the Northern Boreal zone. P. lycopodioides reaches north to Iceland and S Norway in the west and approxi mately to the Middle Boreal zone in the east. P. brevifolium occurs only in the arctic areas. P. turgescens is found in the Scandinavian mountain range, north-eastwards to north-westernmost Finland and to Troms in Norway, and occurs also in (probably) relict localities around the Baltic Sea. P. trifarium occurs throughout the area. The taxonomic position of the genus is discussed.

L. Heden?s, Dept of Botany, Univ. of Stockholm, S-106 91 Stockholm, Sweden

(present address: Dept of Cryptogamic Botany, Swedish Museum of Natural History, S-104 05 Stockholm, Sweden).

Poa Pseudocalliergon (Limpr.) Loeske nepecMOTpeH othoc CeBepHOM EBponu. IlHTepnpeTauHJi pOAa: oh coctomt m3 P. angustifolium Heden?s, hob. BHA, P. lycopodioides (Brid.) Heden?s, hob. kom6.,P. brevifolium

(Lindb.) Heden?s, hob. kom6., P. turgescens (T. Jens.) Loeske h P.

trifarium (Web. & Mohr) Loeske. Poa onpeAejweTca, cp. np., tm?imhhbim

mseTOM m 6jiecKOM BHAa, crpyicrypoH na3yiiiHOH rpynnu kjictok, crpyicr ypoM h uBeTOM oceBbix BOJioc h OKOJiorneH BHAa. Bee bhau, 3a hckjhohc uneM P. trifarium, pacryT b 6oraTux M3BecTbio v3Khx h nacro mcjiko

BOAHbix, nepnoAHMecKH Aaace cyxnx aaGojioneHUbix Mecrax o6nTannfl

(HuorAa Mecra oGHTaima 3ajmBaioTOi +/- 6oraTo?? H3Becrbio boao??). B Ce?epHOH E?pone P. angustifolium pacr?T Ha CKaiiAHHaBCKOM ropiiOM KtaccHBe h b 6ojiee boctohhux TeppnTopnax, a TamKe b CeBepuo?? 6opea jii>hoh 30He. P. lycopodioides AOCTHraeT Ha ceBepe McjiaHAHH, Ha 3anaAe IO^iioh HopBerHH h npMMepiio CpeAHen 6opeajn>HOH 30iibi Ha boctokc

P. brevifolium BcrpenaeTca jiHiiib b apKTHnecKHx TeppHxopH?X. P. tur

gescens BcrpeiaeTCJi tia CKaiiAHHaBCKOM ropiiOM MaccMBe, Ha cesepo BocroKe h ce?epo-3anaAe Ohhji?ihahh h b o6jiacTH TpoMC b HopBernn, oh BCTpenaeTCii (BepO?THo) Tamse b pejiMKTOBux Mecrax BOKpyr BajiTH ??cKoro Mop)i. P. trifarium BCTpenaeTCfl noBCWAy b AanHOM pernone.

06cy;KAaeTC? TaKCOHOMHnecKoe nojio^enwe poAa.

The section Pseudocalliergon of the genus Hypnum was erected by Limpricht (1989) for the species Loeskyp num badium (Hartm.) Paul, Calliergon trifarium (Web. & Mohr) Kindb. and Pseudocalliergon turgescens (T. Jens.) Loeske. Loeske (1907) used the name Pseudocal

liergon at the generic level for C. trifarium, P. turgescens and Campylium longicuspis (Lindb. & H. Arn.) Hede n?d. Paul (1924) treated these species, except L. ba

Accepted 26 August 1991

? LINDBERGIA

80

dium, in Scorpidium, where the species Drepanocladus

lycopodioides (Brid.) Warnst., D. brevifolius (Lindb.) Warnst., D. latifolius (Lindb. & H. Arn.) Warnst, and

Scorpidium scorpioides (Hedw.) Limpr. were also in

cluded. Except for Campy Hum longicuspis, Tuomikoski and Koponen (1979) included the same species in Scor

pidium and added the species Hamatocaulis vernicosus

(Mitt.) Heden?s and H. lapponicus (Norrl.) Heden?s to

LINDBERGIA 16:3(1990)



this genus. While most authors of today (e.g., Smith

1978, Crum and Anderson 1981) do not recognize any

taxon with the name Pseudocalliergon, Nyholm (1965) treats Calliergon trifarium in Calliergon sect. Pseudocal

liergon (Limpr.) Broth, and Karczmarz (1971) treats C.

trifarium and P. turgescens in the same section. Smir

nova (1962) treats Drepanocladus lycopodioides, D.

brevifolius and D. latifolius in Drepanocladus sect. Tur

gidus. In a revision of Scorpidium (s.l.) and Limprichtia,

Heden?s (1989c) suggested that Calliergon trifarium, Pseudocalliergon turgescens and the three Drepanocla

dus species mentioned above are closely related and

that they should be treated in the same genus, Pseudo

calliergon. In the present paper the genus Pseudocal

liergon is taxonomically revised for nothern Europe.

The area covered is the same as treated by Nyholm

(1987).

Material and methods

This revision is based mainly on herbarium material,

but all species except Drepanocladus brevifolius (incl. D. latifolius) were studied in the field as well. All to

gether, more than 1600 specimens of the species recog

nized in Pseudocalliergon were studied. The specimens are located in C, GB, H, ICEL, LD, O, OULU, S,

TRH, TUR, UME and UPS. A few specimens were

also borrowed from LE and from herb. K. Dierssen.

Type material was requested from the following herb

aria (in addition to some of those just mentioned): B, BM, BP, BR, C, FH, G, GL, HBG, JE, KRA, L, LY,

M, MICH, NY, PC, POZ and W. I tried to check the types for names of species, sub

species and varieties suspected to belong to Pseudocal

liergon and reported from Europe. In addition, I tried to check the types of extraeuropean taxa which were

suspected (from information in the protologue or from

earlier synonymizations) to be of importance for the

nomenclature of European species. As regards taxa de

scribed by Sanio, I have mostly followed the judge ments of taxonomic level in Wijk et al. (1964).

From now on I will use the nomenclature suggested in

this paper within Pseudocalliergon to facilitate the un

derstanding of the discussion. Names of vegetation zones follow Ahti et al. (1968: Fig. 9).

Taxonomically important characters

Stem. The stem is unbranched or slightly and irregularly branched in Pseudocalliergon trifarium, slightly and ir

regularly branched in P. turgescens and usually in P.

brevifolium and slightly to more distinctly, but usually irreguarly, pinnately branched in P. lycopodioides and

LINDBERGIA 16:3 (1990)

P. angustifolium. The stem has got a central strand and

the cortex consists of l-3(?4) layers of incrassate cells

(Figs IL, 3L, 5M, 6K, 8L). Stem leaves. The measurements and cross-sections of

the stem leaves were made as described in Heden?s

(1989a, c). In the species with straight, rounded or

apiculate leaves (Pseudocalliergon trifarium, P. turges

cens) both leaf length and width were measured. In the

species with falcate leaves which gradually or ? sud

denly narrow to an acuminate apex (P. augustifolium, P.

lycopodioides, P. brevifolium) only leaf width was mea sured. In all species except P. angustifolium (Fig. 1C) the leaves are normally very broad in relation to their

length (Figs 3C, 5C, D, 6C, 8C) and more or less

strongly concave. These features, together, with the

rather crowded insertion of the leaves, often make the

shoots turgid (again with the exception of P. angustifo

lium). The leaves of species of Pseudocalliergon have

sometimes (P. trifarium) or often a characteristic golden

gloss when dry. This is not found in other species of the

Calliergon-Scorpidium-Drepanocladus (CSD) complex. In addition, the Pseudocalliergon species are often de

veloping a distinct brownish yellow colour which is

rarely parallelled in other CSD complex species (most close are some phenotypes of Drepanocladus sendtneri

(Schimp. ex H. M?ll.) Warnst.). In all five species, the alar cells and those just above

them, the supra-alar cells, are indistinctly delimited

from each other and from the surrounding cells (Figs.

U, 3J, 5K, 6J, 8J). The alar cells are quadrate, rectan

gular or longly rectangular and usually more or less

inflated. The walls may be thin but are mostly thick,

yellow and ?porose. In most specimens the alar cells

form an indistinct, transversely triangular group, but

sometimes they form a single row of cells or are appar

ently undifferentiated. The supra-alar cells are rectan

gular, qradrate or transversely rectangular and are

found in one or several rows along the leaf margin above the alar cells. Sometimes the supra-alar cells are

apparently not differentiated. The ontogeny of the alar

cells is of a modified Drepanocladus aduncus type (He den?s 1987a).

The leaf margin is distinctly denticulate (at least

partly) in Pseudocalliergon angustifolium, smooth or

often partly finely denticulate in P. lycopodioides and smooth or sometimes very finely denticulate in P. brevi

folium and P. turgescens. In P. trifarium the leaf margin is ?smooth.

Axillary hairs. In the genus Pseudocalliergon the axil

lary hairs have 1?2(?3) upper cells. The apical cells tend to be long or very long in comparison with most

other taxa of pleurocarpous mosses (Heden?s 1989d).

There is some variation in this last feature (cf. Figs IM, 3M, 5N, 6L, 8M), but in most specimens the majority of the hairs have got long apical cells. The cell walls of the

upper cells of the axillary hairs are getting yellowish or brownish early, i.e. close to the shoot apex and ? as

soon as they have their mature shape. I have only stud

81



ied this feature in herbarium specimens and not in fresh

material. The colouration may thus be due to prolonged

drying of the material, but, in any case, most other

pleurocarpous species do not have early coloured walls

of the upper axillary hair cells even when studied in

herbarium specimens. Perichaetial leaves. I have not seen mature perichae

tia in Pseudocalliergon angustifolium. In the other spe

cies the inner perichaetical leaves are concave and pli cate. In P. lycopodioides they are ?ovate and gradually or suddenly narrowed to an acuminate point (Fig. 3N). In P. brevifolium the perichaetial leaves are rather simi

lar, but slightly more shortly acuminate (Fig. 50). In P.

turgescens these leaves are even more shortly acuminate

(Fig. 6M) and in P. trifarium they are ?suddenly nar

rowed to an acute or obtuse apex (Fig. 8N).

Sexuality. All species of Pseudocalliergon are dioi

cous.

Features of the proximal branch leaves and of the

rhizoids seem to be of little value in the taxomony of

these species. However, rhizoids are rare and I have not

seen them in Pseudocalliergon brevifolium and P. tur

gescens.

Capsule. I have not seen sporophytes in Pseudocal

liergon angustifolium. In the other species, the capsule is of Amblystegium type (cf. Heden?s 1987b, 1989b). It is approximately horizontal in P. lycopodioides (Fig. 30) and P. brevifolium (Fig. 5P) whereas it is horizontal to inclined in P. turgescens (Fig. 6N) and P. trifarium (Fig, 80). It should be noted that I have only seen one

mature capsule of P. brevifolium. The exostome, and

especially the endostome processes are slightly nar

rowed in P. turgescens. and P. trifarium (as compared with the other species; Figs 6R, 8S), a condition which is most probably due to the often inclined capsules in

these species. The exothecial cells were studied in the

position indicated by Heden?s (1989a, c). Other features of the sporophyte were not found to

be of great value in separating the species.

Ecology. In northern Europe, all species of the genus

Psedocalliergon except P. trifarium tend to grow most

commonly in shallow, lime-rich wetland habitats. The

habitats are often small or very small in size (sometimes

only a few m2) and they commonly dry out perodically, or sometimes the habitats are flushed with ? lime-rich

water. Although P. trifarium differs somewhat from the

other species in its habitat preferences, it is still a spe cies of calcium-rich, but usually deeper and larger wet

lands and it is often growing close to the surface of the habitat. The genus is thus rather homogeneous as re

gards at least some aspects of the habitat preferences of

the included species.

82

Delimitation and phylogenetic considerations

As discussed by Heden?s (1989b, c; see also Heden?s

1988), it seems unlikely that the species of Pseudocal

liergon are closely related to the Warnstorfia-Calliergon

species of Tuomikoski and Koponen (1979) or to the other species which have been included in Scorpidium at one time or another (i.e. those of Scorpidium and

Hamatocaulis as interpreted by Heden?s (1989c) or

Campylium longicuspis (Heden?s 1988)). Characters

supporting a treatment of the Pseudocalliergon species in a different taxonomic position than the Scorpidium and Hamatocaulis species (as interpreted by Heden?s

1989c) are: (1) The absence of red colours, but, instead,

the presence of a characteristic brownish yellow or yel low-brown colour, often with a distinct golden gloss in

Pseudocalliergon; (2) The differences in the structure of the alar cell group; (3) The different kinds of leaf curva ture in Pseudocalliergon, Scorpidium and Hamatocaulis

(in species with curved leaves); (4) The axillary hairs, which have l-2(?3), often yellowish or brownish upper cells in Pseudocalliergon and 2-9 (?11) hyaline upper cells in Scorpidium and Hamatocaulis; (5) The well

developed cortex of incrassate cells, without hyaloder mis in the stems of Pseudocalliergon species as com

pared with the well developed cortex with hyalodermis in Scorpidium species and the weakly developed cortex

without hyalodermis in those of Hamatocaulis; (6) The lower outer peristomial layer is basically cross-striolate

(with occasional irregularities) in Pseudocalliergon whereas it is commonly or always (depending on which

species is considered) dotted or dotted-striate (partly or

entirely) in Scorpidium and Hamatocaulis; (7) The

largest spores tend to be somewhat smaller in Pseudo

calliergon (spore size range: 10.2-17.8 (-19.0) urn) than in Scorpidium (12.0-21.0 urn)) or Hamatocaulis

(10.5-24.5 pui). Features which support the treatment of the Pseudo

calliergon species in the same genus are: (1) The charac

teristic colouration and gloss; (2) The typical structure of the alar cell group (to me it seems very unlikely that

as complicated structures as the alar cell groups should

have developed as similar as in the Pseudocalliergon

species due to chance only); (3) The characteristic axil

lary hairs; (4) The similarities in habitat preferences. It is quite clear that the Pseusocalliergon species be

long to "group IV" (including the Amblystegiaceae, parts of the Hypnaceae and parts of the Leskeaceae/

Thuidiaceae) of Heden?s (1989b), but the exact posi tion within this group is difficult to assess. One may speculate on a relationship between the species of Pseu

docalliergon and those of Drepanocladus s. str. (i.e. the

species around D. aduncus (Hedw.) Warnst.), but at

present I am not aware of any synapomorphy joining

Pseudocalliergon with Drepanocladus s. str. with cer

tainty. The axillary hairs have 1-2 (-3) upper cells in both genera, but this character state is found also in




other taxa usually referred to the Amblystegiaceae (He

den?s 1989d). Features in which Drepanocladus species

differ from those of Pseudocalliergon are: (1) The alar cells are different (usually distinctly more inflated and

more distinctly delimited in Drepanocladus than in

Pseudocalliergon) and their ontogeny is not quite simi

lar in the two genera (Heden?s 1987a); (2) The two

genera differ in the coloration of the plants. The Drepa

nocladus species have not got the golden gloss of the

Pseudocalliergon species and (with the exception of D.

sendtneri) are rarely having brownish yellow or yellow brown colours; (3) The axillary hairs are hyaline and have usually not got markedly elongate cells in Drepa

nocladus; (A) The Drepanocladus species are often

growing in nutrient-rich habitats (rich in nitrogen/phos

phorus) whereas their need for calcium seems to be less

expressed than in the Pseudocalliergon species (again with the exception of Drepanocladus sendtneri, which

seems to be almost as extreme as the Pseudocalliergon

species as regards the calcium demands).

Since I have not been able to find a reasonably prob

able sister group of the Pseudocalliergon species, any

phylogenetic consideration is very uncertain. However,

intuitively one would guess that P. angustifolium, which

is most similar to most species of e.g. Drepanocladus s.

str., Warnstorfia and Campy Hum in leaf shape, is the

more primitive of the Pseudocalliergon species treated

here, and that the broader (P. lycopodioides) and later shorter (P. brevifolium), rounded-apiculate (P. turges

cens) and at last entirely rounded (P. trifarium) leaves have evolved later, as adaptations to different habitats.

In parallel, the capsules have envolved from horizontal

and with more perfect peristomes (P. lycopodioides, P.

brevifolius) to often inclined and with a tendency to

have slightly reduced peristomes (P. turgescens and P.

trifarium). It seems less likely that the ovate leaves,

gradually narrowed to an acuminate apex, should have

evolved from broadly ovate rounded leaves, and that

the perfect peristomes should have developed from the less perfect ones, than that the opposite should have

happened. Although speculative, I do not think that it is

possible to get a more firmly based hypothesis on the

relationships between the Pseudocalliergon species at

present.

Taxonomy

Pseudocalliergon (Limpr.) Loeske

Hedwigia 46: 311. 1907. -

Hypnum sect. Pseudocal

liergon Limpr., Laubm. Deutschl. 3: 547. 1899. -

Dre

panocladus sect. Pseudocalliergon (Limpr.) Broth.,

Nat. Pfl. 1(3): 1035. 1908. - Calliergon sect. Pseudocal

liergon (Limpr.) Broth., Laubm. Fennosk. 486. 1923. -

Lectotype (nov.): Pseudocalliergon turgescens (T. Jens.) Loeske, Hedwigia 46: 311. 1907.

- Hypnum tur


gescens T. Jens., Vid. Medd. Naturh. For. Kj?benh. 1858 (\-A)\ 63. 1858.

Calliergon sect. Trifaria C. Jens., Bot. Not. 1921: 30.

1921. - Lectotype (nov.): Calliergon trifarium (Web. &

Mohr) Kindb., Canad. Rec. Sc. 6(2): 72. 1894. -

Hyp num trifarium Web. & Mohr, Naturh. Reise Schwedens

177. 2f 2a-d. 1804.

Drepanocladus sect. Turgidus Z. Smirn., Not. Syst.

Sect. Crypt. Inst. Bot. Komarov. Ac. Se. USSR 15: 181.

1962. - Lectotype (nov.): Drepanocladus lycopodioides

(Brid.) Warnst., Beih. Bot. Centralbl. 14: 401, 413.

1903. -

Hypnum lycopodioides Brid., Spec. Musc. 2:

227. 1812.

Dioicous. Plants pleurocarpous, medium-sized to ro

bust, sometimes turgid, green, brown-green, yellowish or yellow-brown, often with golden gloss when dry, unbranched to irregularly pinnately branched ?in one

plane. Stem leaves slightly to strongly concave, with

ovate-lanceolate to broadly ovate base, at apex broadly

rounded, suddenly narrowed to apiculate point or grad

ually to ?suddenly narrowed to shortly or longly acum

inate point, straight, and then mostly imbricate, or fal

cate, not plicate, nondecurrent or longly decurrent;

margin smooth to distinctly denticulate, plane; nerve

double and short or single and long, sometimes varying in the same species, in cross-section (when single) bi

convex or plano-convex; cells in mid-leaf linear, thin

walled and eporose or incrassate and por?se, epapil

lose, basal cells wider and sometimes shorter, more

strongly incrassate and more strongly por?se, cells near

leaf apex epapillose; alar cells inflated, slightly inflated or (rarely) hardly inflated, thin-walled or incrassate to

strongly incrassate and then often somewhat por?se,

with yellow walls when mature, in approximately trans

versely triangular group or sometimes in single trans

verse basal row or apparently undifferentiated, always

indistinctly delimited from surrounding cells, ontogeny of modified aduncus type, supra-alar cells in one to

several rows along leaf margin above alar cells, some

times apparently lacking or very indistinct, indistinctly delimited from surrounding cells; branch leaves often

slightly smaller and sometimes narrower than the stem

leaves, proximal branch leaves broad, with broadly rounded or apiculate point, sometimes acuminate.

Pseudoparaphyllia foli?se, broad. Paraphyllia lacking. Rhizoids rare, smooth, reddish brown, slightly

branched and inserted on the stem at or just below the leaf insertion. Stem with central strand (sometimes nar

row) and a usually well developed cortex of thick-walled cells. Axillary hairs with 1?2(?3) upper, early yellowish or brownish cells, well developed and usually with long apical cell, abundant. Inner perichaetial leaves straight

and erect, ovate, plicate; margin plane, not or indis

tinctly bordered at shoulder, smooth or partly dentic

ulate below, more strongly denticulate or with single

83



3

Fig. 1. Pseudocalliergon angustifolium (holotype, S).

A: Shoot. B: Shoot apex. C: Stem leaves. D: Branch leaves. E: Proximal branch

^ leaves. F: Pseudopara phyllium. G: Mid-leaf cells

(of different stem leaves). H: Basal cells near nerve.

I: Leaf margin (mid-leaf). J: Alar cells. K: Cross sections of stem leaves. L:

Cross-section of stem. M:

Axillary hairs. - Scales: a: A. b: B. c: C, D, E. d: F, K. e: G, H, I, J, L, M.

teeth at shoulder; nerve single or branched, ending in

mid-leaf or above; cells smooth; vaginula hairy. Perigo nial leaves from broad basal part suddenly or gradually narrowed to broadly acute or acuminate apex. Calyptra

smooth, cucullate, naked. Seta long, smooth, when dry

dextrorsely twisted below and sinistrorsely twisted

above, sometimes untwisted below or above or dextror

sely twisted above, with central strand and a cortex of

1-3 layers of incrassate cells. Capsule cylindrical,

curved, horizontal or inclined: lid conical; exothecial

84

cells quadrate to longly rectangular, thin-walled or in

crassate (especially longitudinal walls), just below mouth (1?)2-6 rows of isodiametric or transversely

rectangular cells; stomata long-pored, near base of cap

sule; annulus separating. Exostome brownish yellow, well developed; outer peristomial layer in lower part

normally cross-striolate, in upper part papillose and

weakly dentate; transverse ridges of primary peristo mial layer normally developed above; border widened

at zone of transition in outer peristomial layer pattern.




Endostome well developed or with somewhat narrow

processes, yellowish or brownish, with high basal mem

brane, processes not or narrowly perforate, cilia 1-4,

well developed, nodose. Spores finely papillose, mature

in summer.

Chromosome numbers (known only for P. turgescens): n

= 11, 11 + lace. (Fritsch 1982).

Key to the species in northern Europe 1. Stem leaves falcate, from basal leaf portion gradually

or ? suddenly narrowed to shortly to longly acum

inate apex. 2 -

Stem leaves ?straight, from basal leaf portion sud

denly narrowed to apiculate point or leaf apex

broadly rounded. 4

2. Southern or mountainous species. Stem leaves long and longly and sometimes narrowly acuminate (Figs

1C, 3C); nerve single, 3-5(?6)-stratose and (31.5?)

38.5-90(-100) um wide near base. 3 -

Arctic species. Stem leaves comparatively short and

more shortly acuminate (Fig. 5C, D); nerve single

(sometimes branched) or double, when single 2-3

stratose and 21?53 urn wide near base.

. 3. P. brevifolium 3. Medium-sized, not turgid, mainly mountainous spe

cies (distributed in the mountains and in the North ern Boreal zone). Stem leaves narrow at base (Fig.

1C), (0.45)0.56-1.12(-1.23) mm wide at widest

part; margin partly denticulate or finely denticulate

. I. P. angustifolium -

Usually robust, somewhat turgid, southern species

(approximately reaching the Middle Boreal zone). Stem leaves broad at base (Fig. 3C), (0.68-)0.90 1.68 mm wide at widest part; margin sometimes

partly finely denticulate. 2. P. lycopodioides 4. Stem leaves apiculate.4. P. turgescens -

Stem leaves with broadly rounded apex. . 5. P. trifarium

1. Pseudocalliergon augustifolium Heden?s, sp. nov. (Fig. 1)

P. lycopodioidi (Brid.) Heden?s affinis sed debilior, foliis caulis angustioribus et marginibus folii magis valde

denticulatis.

Etymology: The specific epithet refers to the stem

leaves, which are narrower (in relation to their length), than in the other species of the genus.

Holotype: "Sweden: J?mtland, Frostviken, the valley between Mt. Gerven?kko's E and W peaks, 14 Aug 1988, Late snow-bed tow. SW. C. 980m a.s.l., L. Hede

n?s (J88-559)", in S!, isotypes in FH!, LE!.


Rather than raising the varietal name abbreviatus,

which may cause confusion with Drepanocladus abbre

viatus Card, et Broth., to species level, the species is

here described as new.

Synonyms

Drepanocladus sendneri f. pseudorevolvens Wint. &

Monk, in Wint., Hedwigia 49: 384: 1910. - Isotypes:

"Hypnum Sendtneri forma pseudorevolvens mit revolv

ens, Verte! [--] Central-Norwegen, Dovrefjeld, Jerkin,

im Hochmoor an der [--] nach Kongsvold, lOOOMtr.,

August 1908, leg. Dr. Winter", in herb. Winter in JE!;

"Drepanocladus Sendtneri-Wilsoni forma pseudorevol vens Wint. et. M?nkem. [-] Central-Norwegen, Dov

refjeld, Jerkin, 1000M, August 1908, leg, Dr. Winter", in herb. Winter in JE!; another isotype exists in HBG

according to Walther and Martienssen (1976).

Drepanocladus lycopodioides var. abbreviatus Monk.,

Laubm. Eur. 769. 177b. 1927. -

Lectotype (nov.):

"Hypnum lycopodioides Schwgr., fo. (var.) abbreviata

Moenkem. Finmarken, Vads?, in S?mpfen, Juli 1904,

leg. D. Winter, No 3", in HBG!.

Apparently dioicous. Plants green, yellow-brown or

brownish yellow, usually with golden gloss when dry,

medium-sized, not turgid, slightly or more strongly (but

usually irregularly) pinnately branched ?in one plane. Stem leaves slightly concave, not plicate, from erecto

patent to spreading and ovate-lanceolate to rather

broadly ovate base gradually narrowed to falcate or

? strongly falcate and longly and (often narrowly) acuminate, channelled or almost tubular apex, (0.45-)

0.56-1.12(-1.23)mm wide at widest part, not or hardly

decurrent; margin distinctly denticulate or finely dentic

ulate, often partly smooth, plane; nerve single, ending in acumen, (31.5?)38.5-70(?73.5) urn wide near base,

in cross-section plano-convex or biconvex, 3-5-stratose,

with adaxial (thin nerve) or central (thick nerve) cells

widest; mid-leaf cells (28-)31.5-87.5(-110.2) x (3.5-) 4.2?7(?8.5) urn, thin-walled and eporose or sligtly in

crassate and por?se, straight or slightly flexuose, with

longitudinal walls ? parallel except at the cell ends which are square, rounded or shortly fusiformly nar

rowed, basal leaf cells wider and usually shorter, more

strongly incrassate and por?se, cells near leaf apex

smooth; alar cells rectangular, rarely quadrate, not or

usually slightly inflated, thin-walled or incrassate and

then ?porose and with yellow walls when mature, in

approcximately transversely triangular or indistinct

group, indistinctly delimited from surrounding cells, su

pra-alar cells quadrate or shortly rectangular, in 1-7

marginal rows of cells (up to 5(-9) cells long) along leaf

margin above alar cells, indistinctly delimited from sur

rounding cells; branch leaves similar to stem leaves but

85



Fig. 2. Distribution of Pseudocalliergon angustifolium in north ern Europe, based on studied specimens. Open symbols repre sent inexact localities.

narrower, proximal branch leaves broad and rounded,

apiculate or acute to acuminate at apex. Pseudopa

raphyllia foli?se, broad. Rhizoids rare (seen in one

plant), smooth, reddish brown, slightly branched, in

serted at or just below the central parts of the leaf insertion. Stem in cross-section round or flattened, with

central strand and a cortex of 2-3 layers of incrassate

cells. Axillary hairs with 1?2(?3) upper, early yellowish cells, 7-12.5 urn wide, apical cell often long. Mature,

fertilized perichaetia not seen. Perigonial leaves (seen in one plant) from broad base ? suddenly or gradualy

narrowed to acuminate apex; margin smooth or almost

so, distinctly bordered at shoulder. Calyptra and spo

rophyte not seen.

Pseudocalliergon angustifolium is most similar to P. ly

copodioides, but is distinctly weaker and is not turgid, has not narrower leaves (Figs 1C and 3C, respectively) and has got more strongly denticulate leaf margins. The

differences between P. angustifolium and P. brevifolium are discussed under the latter.

Habitat: I have studied Pseudocalliergon angustifolium in the field in the mountains of northernmost J?mtland, central Sweden, during the summers 1988 and 1989. In

this area the species grows in shallow, wet depressions on lime-rich ground, often close to late snow-bed vege tation in the low-alpine region. In the warm summer

1988 some of the depressions with P. angustifolium were almost dry in the middle of August. On the other hand, in the snow-rich year 1989 a large proportion of the

occurrences of the species were still snow-covered in the

beginning of August. I do not know anything about the

86

habitat of the species in the Northern Boreal zone in the eastern part of the area studied.

Distribution: Pseudocalliergon angustifolium is wide

spread in the Scandinavian mountain range and is also

known from scattered localities in the Northern Boreal

zone in more eastern areas (Fig. 2). I have not seen

material from other parts of the world, but it seems

likely that the species occurs also further to the east in

Eurasia. It should be looked for also in northern North

America and in mountain ranges further to the south.

Studied material (excepting types): Sweden: J?mtland, Frostviken, ca. 1 km E of lake at 951 m a.s.l., SE of

Sipmeke, 880-1020 m a.s.l., 13 Aug 1988, L. Heden?s

(J88-473), CANM, G, S. J?mtland, Frostviken, NW

slope of Sipmekjeppe, 880-920 m a.s.l., 13 Aug 1988, L. Heden?s (J88-412), C, S; (J88-438), H, S; (J88 425), S; (J88-436), S; (J88-443), S. J?mtland, Frost

viken, E part of Gerven?kko's top plateau, 1000-1120

m a.s.l., 14 Aug 1988, L. Heden?s (J88-547), S. J?m

tland, Frostviken, S part of Mt. Gerven?kko, 2 Aug 1989, L. Heden?s, S. J?mtland, Frostviken, Jorm,

Brackfj?llet Jul 1926, A. H?lphers, S. ?sele Lappmark, Daunatj?kko, birch reg., 650 m a.s.l., 10 Oct 1931, T.

Arwidson, S. ?sele Lappmark, Dorotea, N. Borgaf

jallen, Genjegetjene, 10 Aug. 1922, C. Stenholm, S.

Lycksele Lappmark, T?rna, Strismasund, 10 Jul 1946, A. H?lphers, S, UME. Lycksele Lappmark, T?rna,

Stora Umevatten, Langfj?llb?cken (251), among dry Salix shrubberies, 16 Jul 1963, O. M?rtensson and G.

Wass?n, UPS. Lycksele Lappmark, Tarna, Abelvattnet,

reg. subalp., the strait betw. the lakes, 15-18 Aug 1967,

O. M?rtensson, UPS. Lule Lappmark, Jokkmokk, Ro

venjaure, 13 Jul 1937, A. H?lphers, S. Lule Lappmark,

Jokkmokk, R?vejaure, 12 Jul 1937, A. H?lphers, S. Lule Lappmark, Jokkmokk, regio Sarjekensis, Unna

Rissavare, vr., 9 Aug 1902, C. Jensen and H. W. Ar

nell, S. Lule Lappmark, G?llivare, Mt. Pierka, 750-945

m a.s.l., 23 Jul 1934, T. G. Halle, S. Torne Lappmark,

Jukkasj?rvi, Abiskojokk, in paludib. reg. betul., 17 Jul

1917, E. J?derholm, HBG, S, UPS. Torne Lappmark,

Jukkasj?rvi, ad lacum Abiskojaure, reg. betul., 8 Aug

1917, E. J?derholm, S, UPS. Torne Lappmark, Jukkas

j?rvi, Abisko Nat. P., P?t j ovare, along "j?kk" from NW to Lake Abiskojaure, reg. alp., 23 Jul 1945, H. Persson

and O. M?rtensson, S, UPS. Torne Lappmark, Jukkas

j?rvi, Tornetr?sk area, K?rsavagge, S of and close to

Lake "681", 16 Aug 1951, O. M?rtensson, UPS. Torne

Lappmark, Jukkasj?rvi, Abisko Nat. P., 1944, G. E.

DuRietz, UPS. Torne Lappmark, Jukkasj?rvi, P?sist

j?rok, reg. alp., 31 Jul 1914, E. J?derholm, UPS. Nor

way: S?r-Tr?ndelag, Dovre, Knudsh?e, 29. 7. 1879, N.

C. Kindberg, S. S?r-Tr?ndelag, Kongsvold/Dovre, n?rdl. Knuds?e, 13-1400 m a.s.l., Aug 1908, Winter,

JE. S?r-Tr?ndelag, Kongsvold, Knudsh?e, Aug 1910, Winter, JE. Finmark, Vads?, Jul 1904, Winter, No. 2,

HBG (syntype of Drepanocladus lycopodioides var. ab




Fig. 3. Pseudocalliergon lycopodioides (Sweden:

Uppland, Edbo, 10 Jul 1867, H. Mos?n, S). A: Shoot. B. Shoot apex. C: Stem leaves. D: Branch leaf. E: Proximal branch leaves. F:

Pseudoparaphyllia. G: Mid-leaf cells (of different

stem leaves). H: Basal cells near nerve. I: Leaf margin (mid-leaf). J: Alar cells. K:

Cross-section of stem leaf. L: Cross-section of stem. M:

Axillarly hair. N:

Perichaetial leaves. O:

Capsule (moist). P:

Exothecial cells. Q: Stoma. R: Exostome tooth. S: Portion of endostome. -

Scales: a: A. b: B, O. c: C, D, E, N, d: F, K. e: G, H, I

J, L, M, P, O, R, S.

3

breviatus M?nk.). Finland: Kuusamo, unweit der Sta

tion Oulanka, 13 Aug 1971, A. v. H?bschmann, OULU. Lapponia enontekiensis, L?t?seno, Tarpoman

taite, 25 Jul 1966, H. Rovainen, H, OULU, TUR.

Lapponia enontekiensis, Kilpiskoski, 24 Aug 1867, J. P.

Norrlin, H, Lapponia enontekiensis, Kilpisj?rvi, 25

Aug 1867, J. P. Norrlin, H. Lapponia enontekiensis,

Kilpiskoski, 8. 1867, J. P. Norrlin, H. Lapponia kemen

sis, Sodankyl?, Peurasuvanto, 21 Jul 1961, R. Ruuhi

j?rvi, H. Ostrobottnia borealis, Rovaniemi rural

comm., Narkaus, Kalkkimaa, (27?E 73553:4652), 153 m


a.s.l., 22 Aug 1989, T. Ulvinen, OULU. USSR: Kola

peninsula, C part, Chibinsk, Tulijok, 25 Jul 1930, A. A. and M. B. Kortjaginy (No. 137), LE. Lapponia mur

manica, peninsula piscatorum, Bumansfjord, Aug 1885, V. F. Brotherus, H, S. Lapponia tulomensis, Pum

manki, Aug 1885, V. F. Brotherus, H. Lapponia pet

samoensis, Petsamo, Kolttak?ng?s, 11 Jul 1932, H. Wa

ris, TUR. Lapponia petsamoensis, Petsamo, Kolttak?n

g?s, 11 Jul 1932, A. V. Auer, TUR.

87



2. Pseudocalliergon lycopodioides (Brid.) Heden?s, comb. nov. (Fig. 3)

Hypnum lycopodioides Brid., Spec. Musc. 2: 227. 1812.

-Stereodon lycopodioides (Brid.) Brid., Bryol. Univ. 2: 824. 1827.

- Hypnum aduncum var. lycopodioides

(Brid.) Dub., Bot. Gall. 2: 561. 1830 (nom. illeg. incl. var. prior.

= Hypnum aduncum var. rugosum Hook. &

Tayl.; see below). -

Amblystegium lycopodioides

(Brid.) DeNot., Cronac. Briol. Ital. 2: 23. 1867. -Hyp num vernicosum var. lycopodioides (Brid.) Vent.,

Nuov. Giorn. Bot. Ital. 17: 173. 1885. -

Harpidium

lycopodioides (Brid.) Lange & C. Jens., Medd. Groen

land 3: 336. 1887. -

Drepanocladus lycopodioides

(Brid.) Warnst., Beih. Bot. Centralbl. 14: 401, 413.

1903. - Hypnum aduncum ssp. lycopodioides (Brid.)

Ren., Rev. Bryol. 33: 91. 1906. -

Scorpidium lycopo diodes (Brid.) Paul, Bryol. Zeitschr. 1: 154. 1918. -

Lectotype (Heden?s 1989c): "Hypnum rugosum Linn.,

Turner, Smith, Web. et Mohr. -

scorpioides Schultz et

alin. autor., In pratis paludosis-turfosis Megapolitanae

copi?se fructic cum Hypno trifario, 1799 zuerst von mir bei Malchin aufgenommen, Musci Frond, assioc. F. IV.

a. 193. [two rows of text which I cannot decipher] H. Stereodon lycopodioides Bryol. Univ.", in herb. Bridel

in B!.

Hypnum aduncum var. rugosum Hook. & Tayl., Muse.

Brit. 111. 1818. (Hypnum rugosum Sm., Fl. Brit. 3:

1325. 1805. (horn, illeg. - non H. rugosum Hedw.,

Spec. Muse. 923. 1801. = Rhytidium rugosum (Hedw.)

Kindb.)). -

Lectotype (nov.): "H. rugosum Fl. Br. non

Hedw. aquo omnio distinctum est. Ex. Anglia.

Hooker", in herb. Swartz (No. 2447) in S!.

Hypnum lycopodioides var. permagnum Limpr., Laubm. Deutschl. 3: 399. 1898.

- Drepanocladus lyco

podioides var. permagnus (Limpr.) Warnst., Beih. Bot.

Centralbl. 13: 415. 1903. -Lectotype (nov.): "Hypnum

lycopodioides var. permagnum Limpr., 27 Mai 1870,

Auf [uninterpretable] Begnatshauser Torfmoor. Im Sa

lem. Oberbaden. Jack. ", in herb. Jack in G!, probable

isolectotypes in H-SOL!, S!.

Drepanocladus wilsoni var. platyphyllus Roth, Hedwi

gia 48: 158, 6 f 5. 1908. - Isotype: "Drepanocladus

Wilsoni var. platyphyllus Roth. Westfalen: Moorgr?ben bei Rheine, 1908, leg. H. Brockhausen, com. Rth.", in

BM!.

Dioicous. Plants green, brown-green or yellow-brown, often with golden glass when dry, robust, somewhat

turgid, slightly or more strongly, but irregularly pin

nately branched ?in one plane. Stem leaves concave,

not plicate, from erecto-patent to almost spreading and

ovate to very broadly ovate base ? gradually narrowed

to falcate or strongly falcate and (acuminate or) longly acuminate, channeled or almost tubular apex, (0.68-)

88

0.90-1.68mm wide at widest part, not or hardly decur

rent; margin smooth or often partly finely denticulate,

plane; nerve single, ending in acumen, (37.0?)42.0

90.0(?100.0) urn wide near base, in cross-section bicon

vex or almost plano-convex, 3-5(?6)-stratose, with

adaxial (thin nerve) or central (thick nerve) cells widest; mid-leaf cells (27.0-)31.5-166 X 5.2-9 urn, thin-walled

and eporose or incrassate and por?se, straight or some

times slightly flexuose, with longitudinal walls ?parallel except at the cell ends which are fusiformly narrowed, rounded or, more rarely, square, basal leaf cells wider

and usually shorter, more strongly incrassate and po

r?se, cells near leaf apex smooth; alar cells rectangular or shortly rectangular, inflated, slightly or strongly in

crassate, somewhat por?se and with yellow walls when

mature, in approximately transversely triangular group

(rarely apparently undifferentiated), indistinctly delim ited from surrounding cells, supra-alar cells shortly rect

angular, quadrate or transversely rectangular, in one or

a few marginal rows of cells (up to ca. 8 cells long) along leaf margin above alar cells (sometimes apparently un

differentiated), indistinctly delimited from surrounding cells; branch leaves similar to stem leaves but smaller,

proximal branch leaves broad and rounded or apiculate at apex. Pseudoparaphyllia foli?se, broad. Rhizoids

rare, smooth, reddish brown, slightly branched, in

serted at or just below the leaf nerve insertion. Stem in

cross-section round or flattened, with central strand and

cortex of 2-3(?4) layers of incrassate cells. Axillary hairs with l-2(?3) upper, early yellowish cells, 8.8-14

urn wide, apical cell often long. Inner perichaetial leaves 2.44-4.40 mm long, concave, ovate and gradually

or suddenly narrowed to acuminate point, plicate; mar

gin plane, not or indistinctly bordered at shoulder, smooth or denticulate below, more coarsely denticulate

or with single teeth at shoulder; nerve single, ending below acumen; cells smooth; vaginula hairy. Perigonial leaves from broad base gradually or suddenly narrowed

to acuminate apex; margin smooth or partly denticulate

above, bordered at shoulder, sometimes indistinctly so.

Calyptra cucullate, smooth, naked. Seta reddish, long,

dextrorsely twisted below and sinistrorsely twisted

above or dextrorsely twisted throughout when dry,

smooth, with central strand and a cortex of 2-3 layers of

incrassate cells. Capsule cylindrical, curved, horizontal

or almost so; lid conical; exothecial cells 23-97 x 15.3

40.8 urn, quadrate to longly rectangular, incrassate, es

pecially longitudinal walls, just below mouth 3-5 rows of transversely rectangular or isodiametric cells; sto

mata long-pored, near base of capsule; annulus separat

ing, of 2-4 rows of cells. Exostome brownish yellow, well developed; outer peristomial layer cross-striolate

below, papillose and weakly dentate above; transverse

ridges of primary peristomial layer normally developed above; border distinctly widened at zone of transition in outer peristomial layer pattern. Endostome well devel

oped, yellowish, smooth or papillose above, smooth

below, with high basal membrane, processes not or




Fig. 4. Distribution of Pseudocalliergon lycopodioides in north

ern Europe, based on studied specimens. Four localities in

Sweden and five in the province Karelia onegensis (USSR) are

not included on the map since I was unable to localize them.

narrowly perforate, cilia 1-3, well developed, nodose.

Spores (10.5)11.5-17.5(?19.0) urn, finely papillose, mature in summer.

Pseudocalliergon lycopodioides is the most robust of the

Pseudocalliergon species with acuminate leaves. The

most important differences between this species and P.

angustifolium and P. brevifolium are given under the

latter two species.

Habitat: Pseudocalliergon lycopodioides grows mainly

in small lime-rich wetland habitats. It is often found in

shallow depressions on flat limestone rocks (often on

alvar), on sea or lake shores or in man-made hollows

and ditches. The last kind of habitat was probably more common before modern agricultural methods were in

troduced. Most of the habitats are subject to periodic desiccation.

Distribution: Besides two finds in Iceland (Dierssen 1973; vouchers seen by me, herb. Dierssen), Pseudocal

liergon lycopodioides has a markedly southern and east

ern distribution in norther Europe (Fig. 4). In the west,

the species reaches northwards only to the Oslo area in

Norway. Further to the east, the species occurs further

to the north. Thus, in eastern Sweden P. lycopodioides

reaches the province G?strikland and in Finland it oc

curs northwards to the provinces Ostrobottnia borealis,

Lapponia kemensis and Kuusamo. The report of this

species from the Kola peninsula (USSR) by Schljakov and Konstantinova (1982) refers to P. angustifolium.

The species is widely distributed in Europe and has also been reported from the Asiatic parts of USSR


(Nyholm 1965, Wijk et al. 1962) and from North Amer ica (Wijk et al. 1962). However, I have not seen mate

rial from other areas than Europe and Janssens (1983)

only reports Pseudocalliergon brevifolium from North

America. The occurrences outside Europe are in need

of confirmation.

The material chosen as lectotype of Hypnum aduncum

var. rugosum Hook. & Tayl. is the only possible speci men which I have seen. Since Swartz died in 1818 (Veg

ter 1986), the specimen in his herbarium cannot have

been collected by Hooker after the description of the taxon (Hooker and Taylor 1818).

In BP, where Limprichts herbarium should be located

according to Sayre (1977), no type material of Hypnum lycopodioides var. permagnum Limpr. exists. However,

in G, where Jack's (the collector) herbarium is located

two syntypes are present. One of these is selected as

lectotype of the name.

Exiccata specimens studied (totally 475 collections

seen): E. Bauer, Musci europaei exsiccati No. 1413

1417, S. V. F. Brotherus, Bryotheca Fennica No. 189a,

S; 189b, H, S. F. Gravet, Bryotheca B?lgica No. 247, S.

Husnot, Musci Galliae No. 940A, S, Mikutowicz,

Bryotheca B?ltica No. 629 + 629a-g, S. R. Ochyra, Musci Poloniae Exsiccati No. 363, S. Rabenhorst,

Bryotheca Europaea No. 752, 752b. 914, 1200, S. Sil

l?n, Musci Frondosi Scandinaviae Exsiccati No. 160,

GB, UPS. Bryophyta D?nica Exsiccata No. 326,

OULU, S, UPS; 472 OULU. Bryotheca Polonica No.

1216, S. Cryptogamae exsuccatae editae a Museo Hist.

Natur. Vindobonensi No. 4866, S. Flora exsiccata Ba

varica: Bryophyta No. 31, S.

3. Pseudocalliergon brevifolium (Lindb.)

Heden?s, comb. nov. (Fig. 5)

Hypnum brevifolium Lindb., Oefv. K. Vet. Ak. Foerh.

23: 541. 1867. (H. brevifolium Schleich, in Brid., Spec. Muse. 2: 149. 1812 (nom. nud. in synon.

= Hetero

cladium dimorphum (Brid.) B. S. & G., cf. Bridel

(1812))). -

Harpidium brevifolium (Lindb.) Lange & C.

Jens., Medd. Groenland 3: 325. 1887. -

Amblystegium

brevifolium (Lindb.) C. Jens, in Luetken, Dijmphna Togt. Zool. Bot. Udbytte 69. 1887.

- Hypnum lycopo

dioides var. brevifolium (Lindb.) Ren. in Husn., Muse.

Gall. 395. 1894 (horn illeg., non Hypnum lycopodioides var. brevifolium Berggr., K. Svensk. Vet. Ak. Handl.

13: 83. 1875; cf. below). -

Drepanocladus brevifolius

(Lindb.) Warnst., Beih. Bot. Centralbl. 13: 401, 416.

1903. - Scorpidium brevifolium (Lindb.) Paul, Bryol.

Zeitschr. 1: 154. 1918. -Lectotype (nov.): "H. brevifo

lium Lindb., Spitsb. Lomme bay. 1861. Malmgren", in

H-SOL!.

Hypnum lycopodioides var. brevifolium Berggr., K.

89



3

Fig. 5. Pseudocalliergon brevifolium (A-C, E-N: Svalbard: Nordfjorden,

Kapp Wijk, S for Laguna, 15 Aug 1973, A. A. Frisvoll,

TRH. D: Svalbard:

Nordfjorden N0, Kapp Wijk, tett ved Oxaashytta,

30 Jun 1973, A. A. Frisvoll, TRH. O: Greenland:

Kangerdluarssuk Fjord, Esersiutilik, 28 Jul 1956, K. Holmen (16505), S. P, R-U:

isolectotype of Hypnum lycopodioides var.

brevifolium Berggr., UPS.

Q: lectotype of

Amblystegium latifolium Lindb. & H. Arn., S). A: Shoot B: Shoot apex. C, D: Stem leaves from different

plants. E: Branch leaf. F:

Proximal branch leaves. G:

PseudoparaphyIlium. H: Mid-leaf cells (of different

stem leaves). I: Basal cells near nerve. J: Leaf margin (mid-leaf). K: Alar cells. L:

Cross-section of stem leaf. M: Cross-section of stem. N:

Axillary hairs. O: Perichaetial leaves. P:

Capsule (moist). Q: Capsule (moist); probably immature, see text. R: Exothecial cells. S: Stoma. T: Exostome tooth. U: Portion of endostome. - Scales: a: A. b: B, P, Q. c: C, D, E, F, O. d: G, L. e: H, I, J, K, M, N, R, S, T, U.

Svensk. Vet. Ak. Handl. 13: 83. 1875. -

Harpidium

lycopodioides var. brevifolium (Berggr.) Lange & C.

Jens., Medd. Groenland 3: 326. 1887. -

Amblystegium

lycopodioides var. brevifolium (Berggr.) H. Arn., Ark.

Bot. 15(5): 100. 1918. - Drepanocladus lycopodioides

var. brevifolius (Berggr.) Moenk., Laubm. Eur. 769,

1927. -

Lectotype (nov.): "127. Hypnum lycopodioides

Schwaegr. ? brevifolium Berggr. Insulae Spetsber

genses: Adventbay. 1868. S. Berggren", in LD!, isolec

totypes in LD!, NY!, S! UPS(2)!.

Hypnum badium x wilsoni San., Bih. K. Svensk. Vet.

90

Ak. Handl. 10(1): 52. 1885. - Lectotype (nov.): "Hyp

num badium x Wilsoni Sanio, Sanio det. Sibiria: Jeni

sei, Tolstoinos, 70?10' n. lat., 25/8 1876, H. Wilh. Ar

nell", in S!, isolectotype in UPS!.

Hypnum aduncum L. ? molle San. a wilsoni Schimp. xxxxholleri San. xxjeniseiense San., Hedwigia 26: 165.

1887. - Amblystegium latifolium var. jeniseiense (San.)

Lindb. & H. Arn., K. Svensk. Vet. Ak. Handl. 23(10): 121. 1890. - Lectotype (nov.): "Amblystegium latifolium

Lindb. et Arn. var. jeniseiense (Sanio)., Hypnum adun

cum L. ?) molle Sanio a) Wilsoni Schpr., xxHolleri




Sanio, Sanio det., Sibiria: Jenisei, Saostrovskij ostrov,

69?40' n. lat., 10/9 1876, H. Wilh. Arnell", in UPS!,

isolectotypes in S(3)!, UPS!.

Amblystegium latifolium Lindb. & H. Arn., K. Svensk.

Vet. Ak. Handl. 23(10): 120. 1890. -

Drepanocladus

latifolius (Lindb. & H. Arn.) Warst., Beih. Bot. Cen

trabl. 13: 401, 415. 1903. - Hypnum latifolium (Lindb.

& H. Arn.) Bryhn, Rep. 2 Norw. Arct. Exp. Fram

1898-1902, 11: 132. 1902 (hom. illeg., non H. latifolium Tayl., London J. Bot. 7: 196. 1848. =

Ectropothecium

latifolium (Tayl.) Jaeg.; cf. Wijk et al. (1964)). -

Scorpi dium latifolium (Lindb. & H. Arn.) Paul, Bryol. Zeitschr. 1: 154. 1918.

- Lectotype (nov.): "Amblyste

gium latifolium Lindb. et Arn., Hypnum aduncum L. ?) molle Sanio a) Wilsoni Schpr., Sanio det., Sibiria: Jeni

sei, Tolstoinos, 70?10' n. lat., 29. 8. 1876, H. Wilh.

Arnell" in S!, isolectotypes in NY!, S!, UPS!.

Diocous. Plants green, brown-green or yellow-brown, often with golden gloss when dry, medium-sized, often

somewhat turgid, usually slightly and irregularly branched ?in one plane. Stem leaves concave, not pli

cate, from erecto-patent to patent and ovate to very

broadly ovate base gradually or ? suddenly narrowed to

falcate or strongly falcate and (shortly acuminate,) acuminate or longly acuminate, channeled or almost

tubular apex, (0.44-)0.60-1.08 mm wide at widest part, not or hardly decurrent; margin smooth or sometimes

partly very finely denticulate, plane; nerve either single and extending 60-75% way up leaf, 21-53 urn wide near

base, in cross-section biconvex, 2-3-stratose and con

sisting of homogeneous cells or with adaxial cells

widest, or nerve double and extending 30-40(-50)%

way up leaf; mid-leaf cells (17.5-)22-79(-87) x 5-10.2

(-11.5) urn, thin-walled and eporose to strongly in

crassate and por?se, straight or slightly flexuose, with

longitudinal walls ? parallel except at the cell ends which are square, rounded or, more rarely, rather

shortly fusiformly narrowed, basal leaf cells wider and

sometimes shorter, more strongly incrassate and po

r?se, cells near leaf apex smooth; alar cells rectangular or longly rectangular, inflated, slightly or strongly in

crassate, partly por?se and with yellow walls when ma

ture, in apparently transversely triangular group, indis

tinctly delimited from surrounding cells, supra-alar cells

rectangular or quadrate (more rarely transversely rect

angular), in three to several rows of cells (up to ca. 10

cells long) along leaf margin above alar cells, indis

tinctly delimited from surrounding cells; branch leaves similar to stem leaves but smaller and sometimes nar

rower, proximal branch leaves broad and rounded or

apiculate at apex. Pseudoparaphyllia foli?se, broad.

Rhizoids not seen. Stem in cross-section round or flat

tened, with central strand (sometimes narrow) and cor

tex of 1-2 layers of incrassate cells. Axillary hairs with

1?2(?3) upper, early yellowish cells, 6.5-12.7 urn wide,

apical cell often long. Inner perichaetial leaves 2.30


3.80 mm long, concave, ?straight and erect, ovate and

gradually to ? suddenly narrowed to acuminate or

shortly acuminate point, plicate; margin plane, unbor

dered or indistinctly bordered at shoulder, smooth or

indistinctly and irregularly denticulate, sometimes with

single teeth at shoulder; nerve single, ending in mid-leaf

or above; cells smooth, vaginula hairy. Perigonial leaves

from broad base gradually or ? suddenly narrowed to

acuminate point; margin smooth or almost so, indis

tinctly bordered or, more rarely, unbordered. Calyptra

smooth, naked (only young seen). Seta reddish, at least

near base, long, untwisted or dextrorsely twisted below

and sinistrorsely twisted above when dry, smooth, with

central strand and a cortex of 2-3 layers of incrassate

cells. Capsule (one ?mature seen) cylindrical, curved, almost horizontal; lid conical; exothecial cells 42-87.5

(?98) x 12.2-24.5 urn, rectangular, incrassate, just be

low mouth 3-5 rows of isodiametric or transversely rect

angular cells; stomata long-pored, near base of capsule; annulus separating, of 2-3 rows of cells. Exostome

brownish yellow, cross-striolate below, papillose and

weakly dentate above; transverse ridges of primary pe ristomial layer normally developed above; border dis

tinctly widened at zone of transition in outer peristomial

layer pattern. Endostome well developed, yellowish,

finely papillose above, very finely papillose below, with

high basal membrane, processes not or narrowly perfo

rate, cilia 1-4, well developed, nodose. Spores 10.5-14

urn, finely papillose, mature in summer.

Pseudocalliergon brevifolium has the most shortly acuminate leaves of the Pseudocalliergon species with

acuminate leaves. The other nothern species of the ge

nus, P. angustifolium, is easily separated from P. brevi

folium by its more longly and more narrowly acuminate

leaves (cf. Figs 1C and 5C, D), its normally single and

stronger nerve and by its more strongly denticulate leaf

margins. P. brevifolium is often distinctly turgid, this not being the case in P. angustifolium. P. lycopodioides differs from P. brevifolium, i. a., in its wider (0.90-1.68 vs 0.60-1.08 mm wide), more longly acuminate leaves

(cf. Figs 3C and 5C, D, respectively) and in its usually single and stronger nerve. There are also differences

between the three species in the length of the mid-leaf

cells, but the overlap is rather great in this character.

The sporophyte characters described by Lindberg and Arnell (1890) are probably based on poor and imma

tured sporophytes (cf. Fig. 5Q; from the lectotype of

Amblystegium latifolium). Lindberg and Arnell (1890) describe the peristome as imperfect and this agrees well

with what I found in the lectotype of Amblystegium latifolium.

Habitat: As I have not seen Pseudocalliergon brevifo lium in the field, the description of its habitat is based on Arnell and M?rtensson (1959) and Brassard (1971). The species is arctic and grows in calcium-rich habitats,

91



^ AH^'^? ?it

Fig. 6. Pseudocalliergon turgescens (Estonian SSR:

Kingissepa, Saarema, L?o, 13 Jun 1989, L. Heden?s

(E-21), S). A: Shoot. B: Shoot apex. C: Stem leaves.

D: Branch leaf. E: Proximal branch leaves. F:

Pseudoparaphyllia. G: Mid-leaf cells (from different stem leaves). H: Basal cells near nerve. I: Leaf margin (mid-leaf). J: Alar cells. K: Cross-section of stem. L: Axillary hairs.

M: Perichaetial leaves. N:

Capsule (moist). O: Exothecial cells. P: Stoma.

Q: Exostome tooth. R: Portion of endostome. -

Scales: a: A. b: B, N. c: C, D, E, M. d: F. e: G, H, I, J, K, L, O, P, Q, R.

in rich fens, on moist excavated soil and in percolation areas. It is often found together with P. turgescens..

Distribution: Within the studied area, Pseudocalliergon

brevifolium occurs only on Svalbard.

The species distribution is circumarctic and I have

seen material from several localities in the arctic parts of

USSR, Alaska, Canada and Greenland.

It should be noted that Hypnum lycopodioides var.

brevifolium Berggr. is not a nomenclatoric synonym of

H. brevifolium Lindb. The latter taxon is also treated in

92

the paper where Berggren described his taxon (Berg

gren 1875). In LD, where Beggrens herbarium is lo

cated, there exists an envelope with two collections

inside ("a)" and "b)") and with the joint label "Hypnum brevifolium Berggr. nova spec. S. Berggren" (no local

ity or date given on this label). The two collections both

belong to Pseudocalliergon brevifolium, but differ in

appearance and have both been distributed as "Musci

Spetsbergens. exsicc. No. 127". An exsiccate specimen with a portion of the collection which agrees best with

the protologue is here chosen as lectotype of Berggrens

(1875) name.




Selected collections studied (totally 49 collections seen):

Svalbard: Ny-?lesund, 7 Jul 1985, J. Ekman, S. Wijde Bay, Ostfjorden, 8 Aug 1899, T. Wulff, S, UPS. Kings Bay, 1868, S. Berggren, S. Liefde Bay, 1868, S. Berg

gren, S, UPS. Prins Charles Forelands Sund, 1868, S.

Berggren, S, UPS. Nordfjorden, 1868, S. Berggren, S,

UPS. Bear Island, Mount Misery, 1868, S. Berggren,

UPS. Kings Bay Distr., the slope of Zeppelinfjeilet "556" above Ny-?lesund, 22 Jul 1956, S. Arnell and O.

M?rtensson, UPS. Kings Bay Distr., Ny-?lesund, rich

fen beside pools immed. E of Gluudneset, 25 Jul 1956, S. Arnell and O. M?rtensson, UPS. Br?ggerhalv?ya,

ved vestlige Huklagunene, 25 Jul 1974, A. A. Frisvoll, TRH. Kongsfjorden, Ny-?lesund, Storv?hut, 11 Jul

1974, A. A. Frisvoll, TRH. Nordfjorden, Kapp Wijk, S for laguna, 8 Jul 1973, A. A. Frisvoll, TRH. Back

fjorden, Trollkjeldene, 9 Aug 1974, A. A. Frisvoll, TRH. Kapp Linn?, inn for radiostasjonen, 1 Jul 1974,

A. A. Frisvoll, TRH. Kongsfj orden, Ny-?lesund, Sol

vatnet, 7 Jul 1974, A. A. Frisvoll, TRH. Br?ggerhal

v?ya, Kvadehuken inn for Huklagunene (vestligst), 25

Jul 1974, A. A. Frisvoll, TRH. Kongsfjorden, Gluud

neset, dam tett ?st for neset, 16 Jul 1974, A.A. Frisvoll,

TRH. Adventfjorden N, Advent City, strand, 23 Jul

1977, A. A. Frisvoll, TRH. Mitrahalv?ya, p? sletta

mellom Collinsodden og Willeberget, 23 Jul 1974, A.

A. Frisvoll, TRH. Reinsdyrflya, litt vest for Worsley

hamna, 24 Aug 1974, A. A. Frisvoll, TRH. Dicksonf

jorden SO, litt S for Myggdalen p? skr?ningen mot

fjorden, 14 Aug 1973, A. A. Frisvoll, TRH.

4. Pseudocalliergon turgescens (T. Jens.) Loeske

(Fig. 6)

Hedwigia 46: 311. 1907. -

Hypnum turgescens T. Jens.,

Vid. Medd. Naturh. For. Kjoebenh. 1858(1-4): 63.

1858. -

Hypnum molle ssp. turgescens (T. Jens.) C.

Hartm., Handb. Skand. Fl. ed. 9, 2: 6. 1864. -

Ster

eodon turgescens (T. Jens.) Mitt., J. Linn. Soc. Bot. 8:

42. 1865 (according to Wijk et al. 1967, but I could not

find the combination there). -

Amblystegium turgescens

(T. Jens.) Lindb., Musci Scand. 33. 1879. -

Hypnum

aduncum ssp. ? molle var. b) turgescens (T. Jens.) San.,

Bih. K. Svensk. Vet. Ak. Handl. 10(1): 38. 1885. -

Calliergon turgescens (T. Jens.) Kindb., Canad. Rec.

Sc. 6(2): 72. 1894. -

Scorpidium turgescens (T. Jens.)

Loeske, Verh. Bot. Ver. Brandenburg 46: 199. 1905. -

Hypnum molle x turgescens C. Hartm. ex Par., Ind.

Bryol. ed. 2, 3: 102. 1905 (nom. inval. err. pro. H. molle

ssp. turgescens (T. Jens.) C. Hartm.). -

Drepanocladus

turgescens (T. Jens.) Broth., Nat. Pfl. 1(3): 1035. 1908. -

Scorpidium scorpioides var. turgescens (T. Jens.)

Moenk., S?sswasserfl. 154. 1914. -

Lectotype (Heden?s

1989c): "Kongsvold, Dovre, 7/56, Th. Jensen, Hypnum

turgescens Th. Jensen in Vidensk. Medd. [etc.]", in C!,

isolectotype in H!.


Hypnum turgescens var. uliginosum Lindb., Oefv. K.

Vet. Ak. Foerh. 23: 539. 1867. -

Calliergon turgescens var. uliginosum (Lindb.) Warnst., Hedwigia 54: 137.

1913. -

Lectotype (nov.): "Hypnum turgescens var. uli

ginosum (19 Febr. 67), Spitsbergen, ad Treurenberg

bay, 1861, leg, Malmgren", in H-SOL!.

Hypnum turgescens var. tenue Berggr., K. Svensk. Vet.

Ak. Handl. 13(7): 91. 1875. - Amblystegium turgescens

var. tenue (Berggr.) C. Jens., Medd. Groenland 15:

433. 1898. -

Calliergon turgescens var. tenue (Berggr.)

Karcz., Monographiae Botanicae 34: 169. 1971. - Lec

totype (nov.): "151b. Hypnum turgescens Seh. ? tenue.

Beeren Eiland: Nordhamnen, 1868, S. Berggren", in

LD!, isolectotypes in FH!, H!, NY!, S(2)!, UPS!, prob able isolectotypes in LD!, NY!,S!.

f Hygrohypnum szaferi Podp. in Szafr., Starunia (Pol ska Akademja Umiejetnosci, Krakow) 1:10. 6. 1934. -

Probable holotype (slide): "Hygrohypnum szaferi Podp.", in KRA!.

Dioicous. Plants green, brown-green or yellow-brown, often with golden gloss when dry, robust, rarely

weaker, turgid, slightly and irregularly branched ?in one plane; shoot apices often falling of and propagating the species vegetatively. Stem leaves strongly concave,

imbricate or sometimes ?spreading, not plicate, ovate

or broadly ovate, upwards suddenly narrowed to short

apiculus, (0.84-)1.52-2.88 mm long, (0.67-)0.80-1.46 mm wide at widest part, not or hardly decurrent; margin smooth or partly very finely denticulate, plane; nerve

mostly double and extending 20-40% way up leaf, in scattered leaves single (then often ill developed and sometimes branched) and extending 60-65% way up leaf; mid-leaf cells (17.9-)21-107(-117) x (5.2-)6.4 11.5 urn, slightly to strongly incrassate, ?por?se,

straight or slightly flexuose, with longitudinal walls

?parallel except at the cell ends which are square,

rounded or ?shortly fusiformly narrowed, basal leaf

cells wider and often shorter, more strongly incrassate

and por?se, cells near leaf apex smooth; alar cells quad rate to rectangular, slightly inflated, incrassate or

strongly incrassate, slightly por?se and with yellow walls

when mature, in approximately transversely triangular

group, indistinctly delimited from surrounding cells, su

pra-alar cells transversely rectangular, quadrate or

shortly rectangular, sometimes very indistinctly delim

ited from both the alar cells and from the lamina cells further up, sometimes more distinct (but still indis

tinctly delimited from surrounding cells) and then in four to several rows of cells (up to ca. 15 cells long) along leaf margin above alar cells; branch leaves similar

to stem leaves or smaller, proximal branch leaves broad

and rounded or sometimes apiculate at apex. Pseudopa

raphyllia foli?se, broad, rare. Rhizoids not seen. Stem

in cross-section round or flattened, with central strand

93



Fig. 7.Distribution of Pseudocalliergon turgescens in northern

Europe, based on studied spedimens and one literature report

(square; Ahti and Roivainen 1968; not possible to locate in H

according to P. Isoviita in litt 1989). Open symbols represent inexact localities. One locality in the Swedish province Torne

Lappmark is not included on the map since I was unable to

localize it.

(sometimes narrow) and cortex of (1?)2-3 layers of

incrassate cells. Axillary hairs with 1?2(?3) upper,

early yellowish cells, 8.9-14 urn wide, apical cell often

long. Inner perichaetical leaves 2.20-3.80 mm long,

concave, ?straight and erect, ovate and gradually or

rather suddenly narrowed to shortly acuminate point,

plicate; margin plane, indistinctly bordered at shoulder,

smooth or denticulate below, denticulate or coarsely denticulate at shoulder; nerve single or branched, end

ing in mid-leaf or above; cells smooth; vaginula hairy.

Perigonial leaves from broad base ?suddenly narrowed

to apiculate or acuminate point; margin smooth or

slightly denticulate near apex, bordered at shoulder,

sometimes indistinctly so. Calyptra cucullate, smooth,

naked. Seta reddish, long, dextrorsely twisted below

and untwisted or sinistrorsely twisted above when dry,

smooth, with central strand and a cortex of (1?)2-3

layers of incrassate cells. Capsule cylindrical, curved,

horizontal to inclined; lid conical; exothecial cells 20.4

88.0 x (7.6-) 11.5-35.6 urn, (quadrate or) shortly to

longly rectangular, thin-walled or with longitudinal walls incrassate, just below mouth 2-6 rows of isodia

metric or transversely rectangular cells; stomata long

pored, near base of capsule; annulus separating, of 2-3

(-4) rows of cells. Exostome brownish yellow, well

developed; outer peristomial layer cross-striolate below

(in single teeth partly irregularly so or with small por tions (0?10(?25)%) irregularly dotted), finely papillose and weakly dentate above; transverse ridges of primary

peristomial layer normally developed above; border dis

94

tinctly widened at zone of transition in outer peristomial

layer pattern. Endos tome well developed but processes sometimes rather narrow, yellowish or brownish, papil lose above, smooth or finely papillose below, with high

basal membrane, processes narrowly perforate, cilia

1-4, well developed, nodose. Spores 10.2-17.0 urn,

finely papillose, mature in summer.

Pseudocalliergon turgescens is easily known by its rather

robust and usually weakly branched shoots with

straight, imbricate or sometimes slightly spreading, ovate or broadly ovate, apiculate and strongly concave

leaves. The species is sometimes propagated vegeta

tively by means of buds which fall of from the shoots (cf. Miller 1985, and references therein).

Habitat. Pseudocalliergon turgescens grows in lime-rich

areas, and mainly in two kinds of habitats. Firstly, it is

found in small wetland habitats, often in shallow de

pressions in flat limestone rocks (often on alvar in the

southern localities). Secondly, it may be found along rills or on rocks flushed with ?lime-rich water. More

rarely, it grows submerged in small lakes or pools.

Distribution. Pseudocalliergon turgescens is widely dis

tributed in the Scandinavian mountain range, from the

sea level on the Norwegian coast to the alpine region. In

addition, the species occurs in numerous localities

around the Baltic Sea and in the Swedish province V?s

terg?tland (Fig. 7). The southern localities are inter

preted as late-glacial relict occurrences by Albertson

(1940). A striking feature of the species distribution

pattern is its apparent absence from the Norwegian

province Finnmark and from the Kola peninsula in

USSR (cf. Schljakov and Konstantinova 1982), except for one locality reported by Karczmarz (1971; material

not seen by me). Outside the study area, Pseudocalliergon turgescens

occurs in scattered areas throughout the Holartic re

gion, in South America, New Guinea (Crum and An

derson 1981) and in Africa (Karczmarz 1971).

Karczmarz (1971) states that the type of Hypnum tur

gescens var. uliginosum Lindb. was collected in the

Swedish province ?land and that the holotype is in L. This must be due to some mistake since Lindberg (1867) described this taxon on material from Svalbard and

Linberg's herbarium is located in H-SOL.

According to Karczmarz (1971), the holotype o? Hyp num turgescens var. tenue Berggr. exists in LD. How

ever, in LD there is no material which with certaintly can be regarded as the holotype. One specimen from

Berggren's herbarium which could possibly be the holo

type has no indication that it is the same material as is

distributed as "Musci Spetsbergens. exsicc. No. 151b."

Thus, the name must be lectotypified on an exsiccate

specimen and the specimen in LD was chosen.

Wijk et al. (1962) state that Hygrohypnum szaferi




Fig. 8. Pseudocalliergon trifarium (Sweden: Sm?land,

J?nk?ping, Munksj?n, Aug 1905, E. L. Ekman, UPS).

A: Shoot. B. Shoot apices. C: Stem leaves. D: Branch leaves. E: Proximal branch leaves. F:

Pseudoparaphyllia. G. Mid-leaf cells (stem leaf). H: Basal cells near nerve. I: Leaf margin (mid-leaf). J: Alar cells. K: Cross-section of stem leaf. L: Cross section of stem. M: Axillary hairs. N: Perichaetial leaves.

O: Capsules (moist). P: Exothecial cells. Q: Stoma. R: Exostome tooth. S: Portion of endostome. -

Scales: a: A. b: B, O. c: C, D, E, N. d: F, K. e: G, H, I, J, L, M, P, Q, R, S.

3

Podp. is an extant taxon. This is, however, not the case,

but the taxon is included in the present revision (despite that it is a fossil one) to clarify the situation. Since none

of the leaves on the slide in KRA can with certainly be said to be the same as the one illustrated in Szafran

(1934), I regard the KRA slide a probable holotype only.

Exsiccata speciments studied (totally 435 collections

seens): E. Bauer, Musci europaei exsiccati No. 1518?

1521, S. Husnot, Musci Galliae No. 788, S, UPS. Miku


towicz, Bryotheca B?ltica No. 142, 142a-c, S. R.

Ochyra, Musci Poloniae Exsiccati No. 669, S. Raben

horst, Bryoyheca Europaea No. 490, S. Bryophyta Sval

bardensia Exsiccata No. 51, S. Cryptogamae exsiccatae

editae a Museo Hist. Natur. Vindobonensi No. 2599a,

b, S. Flora Exsiccata Austro-Hungarica No. 711, S.

95



Fig. 9. Distribution of Pseudocalliergon trifarium in northern

Europe, based on studied material and on information from the Swedish wetland survey (small dots). Open symbols repre sent inexact localities. Two localities in Sweden (Pite Lapp

mark, Lule Lappmark) and one in USSR (Lapponia murman

ica) are not included on the map since I was unble to localize them.

5. Pseudocalliergon trifarium (Web. a Mohr) Loeske (Fig. 8)

Hedwigia 46: 301.1907. - Hypnum trifarium Web. &

Mohr, Naturh. Reise Schwedens 177. 2f 2a-d. 1804. -

Hypnum stramineum var. ? H. trifarium (Web. &

Mohr) Schwaegr., Sp. Muse. Suppl. 1(2): 212. 1816. -

Hypnum sarmentosum a trifarium (Web. & Mohr)

Wahlenb., Suppl. Fl. Lapponiae 67. 1826. -

Stereodon

trifarium (Web. & Mohr) Brid., Bryol. Univ. 2: 825. 1827. -

Amblystegium trifarium (Web. & Mohr) De

Not., Cronac. Briol. Ital. 2: 23. 1867. -

Calliergon trifa rium (Web. & Mohr) Kindb., Canad. Rec. Sc. 6(2): 72. 1894.

- Drepanocladus trifarius (Web. & Mohr) Broth,

ex Par., Coll. 10. 1909 (according to Wijk et al. 1962; not seen by me).

- Scorpidium trifarium (Web. & Mohr)

Paul, Bryol. Zeitschr. 1: 154. 1918. -

Acrocladium trifa rium (Web. & Mohr) Richs. & Wall., Trans. Brit.

Bryol. Soc. 1(4): 25. 1950. - Lectotype (Heden?s

1989c): "Hypnum trifarium Weber et Mohr. Upsaliae.

[H. Stereodon trifarius. Bryol. Univ.]" in herb. Bridel in

B!, probable isolectotype in C?.

Dioicous. Plants green, brown-green, yellowish or yel

low-brown, medium-sized (rarely almost robust),

slightly turgid, unbranced or slightly and irregularly branched ?in one plane. Stem leaves concave or

strongly concave, imbricate or ?spreading (more

rarely), not plicate, broadly or very broadly ovate with

broadly rounded apex, 1.08-2.24 mm long, 0.80-1.68

96

mm wide at widest part, longly and broadly decurrent

(often about half way down to leaf below); margin smooth or almost so, plane; nerve single, extending

60-90(-95)% way up leaf, 33-68.8 urn wide near base, in cross-section plano-convex, 2-3-stratose and consist

ing of homogeneous cells or adaxial cells widest; mid

leaf cells (30.6-)33.1-104(-107) x 5.1-10.8 urn, in

crassate and por?se or, more rarely, thin-walled and

eporose, straight or slightly flexuose, with longitudinal walls ? parallel except \at the cell ends which are

(square,) rounded or fusiformly narrowed, basal leaf

cells wider and sometimes shorter, more strongly in

crassate and por?se, cells near apex smooth; alar cells

longly rectangular, inflated, incrassate or strongly in

crassate, somewhat por?se and with yellow walls when

mature, in single transverse row or, more commonly, in

approximately transversely triangular group, indis

tinctly delimited from surrounding cells, supra-alar cells

variable, quadrate to rectangular, sometimes forming an indistinct transition from alar cells to lamina cells

further up, sometimes more distinct (but still indis

tinctly delimited from surrounding cells) and then of four to sveral rows of cells (up to ca. 10 cells long) along leaf margin above alar cells; branch leaves somewhat

smaller than the stem leaves, proximal branch leaves

broad and rounded at apex. Pseudoparaphyllia foli?se, broad. Rhizoids rare, smooth, reddish brown, slightly

branched, inserted at or just below the leaf nerve in

sertion. Stem in cross-section round or flattened, with

central strand and cortex of (1?)2-3 layers of incrassate

cells. Axillary hairs with 1-2 upper, early yellowish cells

(rarely hyaline), 8.9-12.8 urn wide, apical cell often

long. Inner perichaetial leaves 2.32-3.20 mm long, con

cave, straigth and erect, ovate, ?suddenly narrowed to

obtuse or acute apex, plicate; margin plane, unbordered

and smooth or irregularly denticulate, especially at

shoulder; nerve single, ending in upper half of leaf; cells

smooth; vaginula hairy. Perigonial leaves from broad

base gradually or, more rarely, ?suddenly narrowed to

broadly acute to acuminate apex; margin smooth or

partly indistinctly denticulate, not or indistinctly bor dered at shoulder. Calyptra cucullate, smooth, naked.

Seta reddish, long, not or dextrorsely twisted below and

sinistrorsely twisted above when dry, smooth, with cen

tral strand and a cortex of l-2(?3) layers of incrassate

cells. Capsule cylindrical, curved, horizontal to inclined; lid conical; exothecial cells 25.5-63.7 x 12.8-28 urn,

(quadrate or) shortly to longly rectangular, with longi tudinal walls somewhat incrassate, just below mouth

(1?)2-5 rows of isodiametric or transversely rectan

gular cells; stomata long-pored, near base of capsule; annulus separating of 2-4 rows of cells. Exostome brow

nish yellow, well developed (sometimes teeth somewhat

narrow); outer peristomial layer cross-striolate below,

papillose and weakly dentate above; transverse ridges of primary peristomial layer normally developed above; border narrow below, distinctly widened at zone of

transition in outer peristomial layer pattern.




Endostome well developed but processess often rather

narrow, yellowish or brownish, papillose above, smooth

or very finely papillose below, with high basal mem

brane, processes not or narrowly perforate, cilia 1-3,

well developed, nodose. Spores 10.4-17.8 urn, finely

papillose, mature in summer.

Pseudocalliergon trifarium is easily known by its un

branched or almost unbranched shoots with straight and

imbricate or more rarely ?spreading, concave and

broadly or very broatly ovate leaves with broadly rounded apices. Sometimes the shoots of P. trifarium are possible to divide into distinct segments, which may correspond with growth seasons. In the lower part of

each segment the leaves are smaller and more strongly imbricate than in the upper portion of the same seg

ment. The transition between the differently sized

leaves is gradually within each segment, whereas the

transition between two segments is rather sudden.

Habitat: Pseudocalliergon trifarium grows in lime-rich,

deep fens. It is often found as single shoots among other

mosses (especially Scorpidium scorpioides). The species occurs from the sea level to the alpine region of the

mountains.

Distribution: Pseudocalliergon trifarium occurs through out the area (Fig. 9), but seems to avoid the coastal

districts in some areas. This may be due to a lack of

suitable habitats. P. trifarium is sometimes considered

to be a rather rare species in northern Europe, but, as

the wetland surveys in parts of northern Sweden have

shown (Fig. 9, small dots) this may be due to lack of

investigations rather than to anything else.

The species has been found in Europe, Asia and

North America. According to Crum and Anderson

(1981) it occurs in Haiti and Venezuela as well.

Exsiccata specimens studied (totally 633 collections

seen): E. Bauer, Musci europaei exsiccati No. 1526, S.

V. F. Brotherus, Bryotheca Fennica No. 83, H, S. V. F.

Brotherus, M. F. E. No. 145, H. Husnot, Musci Galliae

No. 449, 696, S. W. Migula, Kryptogamae Germaniae,

Austriae et Helvetiae exsiccatae No. 175, S. Mikutow

icz, Bryotheca B?ltica No. 144, 144a, S. R. Ochyra, Musci Poloniae Exsiccati No. 169, S. Rabenhorst,

Bryotheca Europaea No. 300, 751, S. Bryotheca Polon

ica No. 86, 896, 1644, S. Flora Exsiccata Austro-Hun

garica No. 1929, S. Flora exsiccata Bavarica: Bryophyta No. 95, S. Hepaticae et Musci URSS Exsiccati No. 50, H, S. Musci Macroregioni Meridionali Poloniae Ex

siccati No. 79, S. Westfalens Laubmoose No. 361, 361b,

S.


Excluded taxa

Hypnum badium Hartm., Handb. Skand. Fl. ed. 5: 332.

1849. -

Drepanocladus badius (Hartm.) Roth, Eur.

Laubm. 2: 569. 1904. -

Loeskypnum badium (Hartm.)

Paul, Bryol. Zeitschr. 1: 155. 1916. -

Lectotype (nov.): "25. Hypnum badium Hn. Lui. 1pm. Ananas. 23.6.48.

HH", in herb. Hartman in UPS!.

In herb. Hartman in UPS there are syntypes from both localities mentioned in the description of Hypnum ba

dium (Hartman 1849). The specimen chosen as lecto

type is the richer one. Hypnum badium was placed in

Hypnum sect. Pseudocalliergon by Limpricht (1899).

Hypnum illecebrum Schultz, Fl. Starg. 318. 1806."

"Hypnum illecebrum Schultz" is stated to be a synonym of Pseudocalliergon trifarium by Karzmarz (1971). However, in Schultz (1806) reference is made to Bridel

(1801: 91), where H. illecebrum Brid. is described. The latter is certainly not a synonym of P. trifarium, but of

Scleropodium tourretii (Brid.) L. Koch (Koch 1949). Thus, Karczmarz's (1971) synonymisation is not cor

rect.

Drepanocladus latinervus Warnst., Beih. Bot. Cen

tralbl. 13: 416. 1903. - Amblystegium latinerve Lindb. in Lindb. & H. Arn., K. Svensk. Vet. Ak. Handl. 23(10): 121. 1890 (nom. nud. in synon.

= A. latifolium var.

jenisseiense (San.) Lindb. & H. Arn.). -

Hypnum lati

nerve H. Arn. in Warnst., Beih. Bot. Centralbl. 13: 416.

1903 (nom. nud. in synonym. =

Drepanocladus latiner

vus Warnst.). -

Holotype: "A. latinerve n. sp., Amblys

tegium n. sp., Sibirien, Jenisei, Tolstoinos, 70?10' n.

lat., 25. 8. 1876, H. With. Arnell", in H-SOL!, isotype in S!.

The type material belongs to Drepanocladus sendtneri

or a taxon close to this species.

Amblystegium longicuspis Lindb. & H. Arn., K.

Svensk. Vet. Ak. Handl. 23(10): 123. 1890. -

Pseudo

calliergon longicuspis (Lindb. & H. Arn.) Loeske, Hed

wigia 46: 311. 1907. - Drepanocladus longicuspis

(Lindb. & H. Arn.) Broth., Nat. Pfl. 1(3): 1035. 1908. -

Scorpidium longicuspis (Lindb. & H. Arn.) Paul, Bryol. Zeitschr. 1: 154. 1918.

- Campylium longicuspis (Lindb.

& H. Arn.) Heden?s, Linbergia 14: 144. 1988 (publ. 1989).

- Lectotype (Heden?s 1988): "Amblystegium lon

gicuspis c. fr.!, Sibiria: Jenisei, Dudinka, 4/8 1876, H.

Wilh. Arnell. Ex HERBARIO S. O. LINDBERGH in S!, isolectotypes in S(3)!, H-SOL, UPS(2)!.

The taxonomic position of Amblystegium longicuspis is discussed by Heden?s (1988).

Hypnum lycopodioides var. genuinum San., Bot. Cen

97



tralbl. 4(Beil. 2): 24. 1880. - Amblystegium lycopo

dioides var. genuinum (San.) Lindb. & H. Arn., K.


Lectotype

(Heden?s 1989c): "83. Hypnum lycopodioides Schw. a

geninum Sanio, Lyck: Rothes Bruch in einigen Torf

l?cher, 5. 5. 1873, C. Sanio", in S!.

Despite the name "genuinum", the type material be

longs to Hamatocaulis vernicosus (Mitt.) Heden?s (He den?s 1989c).

Stereodon vernicosus Mitt., J. Linn. Soc. Bot. 8: 43.

1864. -

Hypnum lycopodioides var. vernicosum (Mitt.)

San., Bot. Centralbl. 4(Beil. 2): 23. 1880. -Hamatocau

lis vernicosus (Mitt.) Heden?s, Lindbergia 15: 27. 1989

(publ. 1990). -

Lectotype (Heden?s 1989c): "Hypnum vernicosum Lindb., c? et c. fr., Sthlm, in palude ad

Carlsbergs park, 20 Julii 1858, leg. S. O. Lindberg", in H-SOL!.

This species is discussed by Heden?s (1989c).

Hypnum sendtneri var. ? wilsonii Schimp., Musci Eur.

Nov. Bryol. Eur. Suppl. fase. 3-4 (Mon.): 3. 3. 1866. -

Hypnum lycopodioides ssp. wilsonii (Schimp.) Ren. in

Husn., Muse. Gall. 375. 107 f. 4a, 5b. 1894.

This is a synonym of Drepanocladus sendtneri (Isoviita and Koponen 1981).

Hypnum lycopodioides ssp. wilsonii var. flageyi Ren. in

Husn., Muse. Gall. 395. 113 f. 3, 4. 1894. -

Lectotype

(nov.): "Hypnum Wilsoni Lindb. var. Flageyi Ren.,

Gallia = Jura: Pontarlier. leg. Flagey", in HBG (ex herb. Renauld)!, isolectotype in S!.

Since no type material seems to exist in PC (herb.

Renauld), a specimen in HBG is chosen as lectotype. The material belongs to Drepanocladus sendtneri.

Hypnum lycopodioides x fluitans San., Bih. K. Svensk.

Vet. Ak. Handl. 10(1): 50. 1885. - Amblystegium lyco

podioides x fluitans (San.) Lindb. & H. Arn., K.


Lectotype

(nov.): "Hypnum lycopodioides x fluitans Sanio, Sanio

det., Sibiria: Jenisei, Fatjanova, 64?5' n. lat., 10. 7.

1876, H. Wilh. Arnell", in S!, isolectotype in S!.

The material belongs to Warnstorfia fluitans (Hedw.) Loeske.

Doubtful taxa and nomina nuda

Hypnum aduncum var. hamatum B. S. & G., Bryol.

Eur. 6: 112. 606 8. (fase. 57-61 Mon 36. 24, 2 e). 1854. -

Hypnum lycopodioides ssp. wilsonii var. hamatum (B.

S. & G.) Ren. in Husn., Muse. Gall. 370. 1894.

98

No certain type material exists. However, in BM (under the name Hypnum lycopodioides) one collection with

the label "Auf Torfmooren u Mecklenburg. Hth Schultz.

Funk. 41." exists. I regard this as a possible isotype of

the name. It belongs to Scorpidium scorpioides (Hedw.) Limpr.

Hypnum aduncum ssp. ? molle var. xxxbinerve San.,

Bih. K. Svensk. Vet. Ak. Handl. 10(1): 38. 1885.

I have not seen original material of this taxon.

Amblystegium badium x latifolium H. Arn. in Lindb. & H. Arn., K. Svensk. Vet. Ak. Handl. 23(10): 122. 1890.

(nom. nud. in synon. =

Hypnum badium x wilsonii

San.; see under Pseudocalliergon brevifolium).

Hypnum calcareum Lindb. in R. Hartm., Bryac. Scand.

Exs. n. 388. 1868 (nom. nud.). -

Hypnum turgescens var. calcareum Podp., Consp. 584. 1954 (nom. nud. in

synon. =

Pseudocalliergon turgescens fo.).

Hypnum lycopodioides var. falcatum Hobk., Syn. Brit.

Moss 169. 1873.


Drepanocladus lycopodioides var. gracilis Jaap. in

Warnst., Krypt. Fl. Brandenburg 2: 1027. 1906 (nom. nud. in synon.

= Pseudocalliergon lycopodioides).

Hypnum lycopodioides x aduncum var. wilsonii San.,

Hedwigia 26: 214. 1887.


Hypnum lycopodioides x fluitans var. exannulatus San.,

Hedwigia 26: 212. 1887.


Hypnum lycopodioides (lapponicum) x fluitans var. ex

annulatum San., Bih. K. Svensk. Vet. Ak. Handl. 10

(1): 50. 1885.


Hypnum lycopodioides x revolvens San., Bot. Cen

trale. 13: 432. 1883.


Hypnum spirale Schleich, in Schwaegr., Spec. Muse.

Suppl. 1(2): 212. 1816 (nom. nud. in synon. = Pseudo

calliergon trifarium).




Calliergon trifarium var. giganteum Warnst., Hedwigia

57: 124. 1915.

I have not seen original material of this taxon. Accord

ing to Prof. H. Hertel (in litt. 1989) no original material exists in herbarium Holler (one collector) in M.

Calliergon turgescens var. patens Karcz., Monographiae

Botanicae 34: 167. 1971. -

Holotype: "GFR, Nieder

bayern, Landau a. I., Kiesgrube bei Griepenau, umweit

Bahnhof Pilstig, 10. XII. 1914, leg. H. Paul", in W (not seen by me; according to Dr. U. Passauer (in litt. 1989) the type material cannot be found in W at present).

Hypnum uliginosum Schleich, in Web. & Mohr, Bot. Taschenb. 319. 1807 (nom. nud. in synon.

= Pseudocal

liergon trifarium).

Acknowledgements - I thank the curators of the herbaria men

tioned in this paper for large loans of material and/or invalu

able help with locating type material. W. R. Buck, R. Grolle, R. Ochyra and the staff at BM were helpful with literature

references not available in Stockholm. M. L?froth was very

helpful with some of the fieldwork and also provided informa

tion from the Swedish wetland survey. I thank T. -B. Engel mark, E. Nyholm and N. Lundqvist for valuable discussions

and help with various problems. Part of the fieldwork was

supported by grants from Karin and Axel Binnings foundation

(the Royal Swedish Academy of Sciences).

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The Genus Pseudocalliergon in Northern Europe