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Some Additions to the Bryophyte Flora of Dominica, West Indies

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Haussknechtia Beiheft 9 (RlCLEF-GROLLE-Festschrift): 317-331; ) 999

ALFONS SCHÄFER-VERWIMP

Some Additions to the Bryophyte Flora of Dominica, West Indies

During a two week stay on the island of Dominica, Lesser Antilles, from 18 to 29 May
1996 the author and his wife have collected 470 specimens of bryophytes among them 32
species which are here firstly reported for the island (indicated by *). Some rarely collec-
ted bryophytes are told with, too. Remarks on ecology äs well äs on the distribution of
several species are given. One species new to science, Bromeliophila helenae, was already
published by GRADSTEIN (1997). Macrocolura is considered äs a mere section (section
Lingua AST) of Colura.
There are no checklists of bryophytes from Dominica; however, many of the mosses are
enumerated by BARTRAM (1955) with additions by TOWNSEND (1994), the pleurocarpous
mosses from the West Indies are treated by BÜCK (1998), and most of the hepatics are
listed by SPRUCE (1895) and FULFORD (1987). Other - often single - records are occurring
in scattered literature. Specimens are deposited in the private herbarium of the author,
duplicates in other herbaria äs indicated.

Hepaticae

Acanthocoleus aberrans (LlNDENB. & GOTT.) KRUUT in Bryophyt. Biblioth. 36: 62. 1988.
(var. aberrans).
= Acanthocoleus fulvus R. M. SCHUST. in Bull. Torrey Bot. Club 97: 339. 1970.
Roseau Valley, on bark of shrubs in very humid low forest around Freshwater Lake, 750 m, c.per.,
18039.
Distr.: A rather rare neotropical taxon which has been gathered only a few times in Mexico, Costa
Rica, Guatemala, Jamaica, French Guiana. From Dominica hitherto known only from the type of
Acanthocoleus fulvus (GRADSTEIN 1994).

Aphanolejeunea angustissima STEPH., Spec. Hep. 5: 859. 1916.

= Aphanolejeunea gracüis AST in Rev. Bryol. Lichenol. 16: 21, Abb. 2. 1947, syn. fide Poes & LÜCKING
1997.
Morne Diablotin, 17931pp, see under Aphanolejeunea costaricensis.
Distr.: Widespread in the Neotropics and known from Colombia, the Guianas, El Salvador, Costa
Rica, Guadeloupe, Galäpagos Islands (LÜCKING 1995, under A. gracilis); Guatemala (type);
Mexico (JOVET-AST 1960), and Brazil (unpublished records by the author); very recently reported
from Dominica by EGGERS (1998), äs A. gracilis.

•'Aphanolejeunea camillü (LEHM.) R. M. SCHUST., Hep. Anthoc. N. Amer. 4: 1297. 1980.

Morne Diablotin, rain forest on westcrn slope, epiphyllous on Selaginella, growing with
Aphanolejeunea sicaefoüa (GOTISCHE) A. EVANS, 620 m alt., 17915, det. T. PÖCS.
Distr.: Widespread in the Neotropics from Brazil and Argentina to Mexico and the West Indies
(Jamaica, Cuba). Seems to be the first record for the Lesser Antilles.
 318

*Aphanolejeunea cingens HERZOG in Svensk Bot. Tidskr. 46: 104. 1952.

Morne Diablotin, rain forest on western slope, epiphyllous, growing with Cyclojeunea peruviana
(LEHM. & LlNDENB.) A. EVANS, Cydolejeunea convexistipa (LEHM. & LlNDENB.) A. EVANS,
Aphanolejeunea costaricensis BERNECKER, Aphanolejeunea cyathiphylla HERZOG, and Crosso-
mitrium patrisiae (BRID.) MÜLL. Hai., 17931pp., det. A. BERNECKER.
Distr.: Ecuador (type), Costa Rica (GRADSTEIN & al. 1994). First record for the West Indies and
new to Dominica where it is probably a rare species, only once collected. This characteristic
species was described and figured by HERZOG (1952: fig. 25a-e) and LÜCKING (1995: Abb. 13).
*Aphanolejeunea costaricensis BERNECKER in Nova Hedwigia 66: 164. 1998.
Morne Diablotin, rain forest on western slope, epiphyllous, associated with Cydolejeunea
peruviana (LEHM. & LlNDENB.) A. EVANS, Aphanolejeunea cyathiphylla HERZOG, A. angustis-
sima STEPH., Odontolejeunea hmulata, and Leptolejeunea radicosa, 850 m, 17931 pp, det. A.
BERNECKER.
Distr.: A. costaricensis was recently described from Costa Rica and hitherto known only from the
type collections. (LÜCKING 1995: Abb. 14; BERNECKER-LÜCKING 1998a: fig. 1).

''Aphanolejeunea cyathiphylla HERZOG in Svensk Bot. Tidskr. 46: 105. 1952.

Roseau Valley, rain forest on the track from Laudat to Middleham Falls, epiphyllous, growing
between Cydolejeunea convexistipa (LEHM. & LlNDENB.) A. EVANS, 60 m alt., 17697, det. A.
BERNECKER; Morne Trois Pitons, elfin forest on northwestern slope, epiphyllous, associated with
Cydolejeunea peruviana (LEHM. & LlNDENB.) A. EVANS and Drepanolejeunea crucianella
(TAYL.) A. EVANS, 1040 m alt., 17840/A, det. A. BERNECKER; Morne Diablotin, 1793 Ipp., see
under Aphanolejeunea costaricensis.
Distr.: Hitherto known only from Ecuador (type) and El Salvador (WINKLER 1967). New to the
West Indies.
Aphanolejeunea sicaefolia (GOTTSCHE) A. EVANS in Bull. Torrey Bot. Club 38: 277. 1911.
Rain forest on Emerald Pool, epiphyllous on fern fronds, growing with Dactylolejeunea
acanthifolia R. M. SCHUST. and numerous other small Lejeuneaceae, 400 m alt., 17737/C; Roseau
Valley, epiphyllous in rain forest on the track between Laudat and Valley of Desolation, with
Trichocolea brevifissa STEPH., 660 m alt., 17856/B; ibid., epiphyllous on fern fronds, together
with Colura tenuicornis (A. EVANS) STEPH., 700 m alt., 17892; Morne Diablotin, rain forest on
western slope, epiphyllous, at 620 m and 800 m alt., 17915pp. (mixed with Aphanolejeunea
camillii), 17927; all det. PÖCS.
Distr.: Widespread in the Neotropics from the West Indies (Puerto Rico, Cuba, Guadeloupe,
Trinidad) to Central and South America (Mexico, Costa Rica, Brazil, the Guianas, Colombia,
Ecuador, and Galäpagos Islands). Probably a rather frequent species on Dominica from where it
was hitherto known only from a „hidden" record by SCHUSTER (1970: 343, mixed with the type of
Dactylolejeunea acanthifolia R. M. SCHUST.).
This species was (described and) figured by STEPHAN! (1888: Tafel XII, figs. 21-24, äs
Cololejeunea sicaefolia), EVANS (1911: pl. 12, figs. 18-26), GRIFFIN (1979: pl. XI, figs. 512-513),
and SCHUSTER (1980: fig. 770:1-2).
ifAphanolejeunea verrucosa AST in Rev. Bryol. Lichenol. 16: 25. 1947.
Roseau Valley, epiphyllous in rain forest on the track between Laudat and Valley of Desolation,
710 m alt., 17891, det. A. BERNECKER & T. PÖCS; ibid., track from Laudat to Freshwater Lake,
culture zone, epiphyllous on Citrus, 700 m alt., 17629, det. T. PÖCS; Morne Diablotin, rain forest
on western slope, epiphyllous, associated with Crossomitrium patrisiae and Taxilejeunea sp.;
870m elev., 17939/B, det. T. PÖCS; rain forest on Emerald Pool, epiphyllous on fern fronds,
400 m alt., 17727/D, det. T. PÖCS.
Distr.: A. verrucosa seems to be widespread but scattered through the Neotropics and is known
from Guadeloupe (type) and Martinique (JOVET-AST 1947: 25, 44), El Salvador (WlNKLER 1967:
 319

316), Mexico (JOVET-AST 1960: 34), and Colombia (WINKLER 1970: 951). First record for
Dominica.
Beside the above mentioned seven species of the genus Aphanolejeunea there is reported A.
microscopica (TAYLOR) A. EVANS var. exigua (A. EVANS) A. LÜCKING & PÖCS (SCHUSTER
1992: 357, äs A. exigua EVANS) and A. longifolia JOVET-AST (SCHUSTER 1992: 355, äs „longi-
folia phenotype", EGGERS 1998) from Dominica. Thus the total number of species in this genus
raised to nine, a high number compared to, for example, Brazil, Costa Rica, and Mexico, each
country with twelve species known, or nine from Cuba, six from El Salvador, and five from
Colombia (PÖCS & LÜCKING 1997). It may be noticed that all records of this frequent but easily
overlooked genus were done not before 1992, except one by SCHUSTER (1970).
Arachniopsis diacantha (MONT.) M. HOWE in Bull. Torrey Bot. Club 29: 288. 1902.
= Arachniopsis coactilis SPRUCH, On Cephalozia: 85. 1882.
Morne Trois Pitons, rain forest on northern slope, at base and roots of big tree destroyed by
cyclone, growing with Cephalozia crassifolia, Zoopsis antillana, and a few plants of Zoopsidella
monodactyla, 550-600 m alt., 17739, conf. VÄNA (PRC), 17740; ibid, elfin forest on northwestern
slope, epiphytic at base of tree fern, c. per., 900 m alt., 17831; Roseau Valley, rain forest on the
track from Laudat to the Valley of Desolation, at base of tree fern, associated with Cephalozia
crassifolia (LlNDENB. & GOTTSCHE) FULFORD and Prionolejeunea spec., 600 m alt., 17850/A;
Morne Diablotin, rain forest on western slope, epiphytic on stump of tree fern; 650 m alt.,
17916/A.
Distr.: A typical Afro-Amcrican species, known from tropical South America (Venezuela,
Colombia, Peru, Brazil), Cuba, Trinidad, in East Africa from Rwanda to Cape, in West Africa
from Sierra Leone to Gabon, äs well äs from Madagascar and the Macarenes (GRADSTEIN & al.
1983, with distr. map, pl. 111/14; distr. map also in GROLLE 1969: fig. 11:3). Surprisingly there is
only one „hidden" record from Dominica by SCHUSTER (1966b: 440, admixed to the type of
Phycolepidozia exigua R. M. SCHUST.); obviously a widespread and rather frequent species but
overlooked by previous collectors due to the small plants growing between other small hepatics,
preferrably at the base of tree ferns or less abundant at the base of old trees and decayed wood,
only rarely in pure mats.
*Cephaloziella antillana (SPRUCE) FULFORD in Mem. New York Bot. Gard. 11(4): 405.
1976.
Roseau Valley, rain forest on Trafalgar Falls, un humid rock, 300 m, 17606/A, det. VÄNA.
Distr.: Seems to be an endemic of the West Indies: Cuba, Jamaica, Puerto Rico, Guadeloupe and
Martinique (FULFORD 1976, with fig. la-p, pl. 110). It could be expected to occur also on
Dominica.
*Cephaloziella granatensis (J. B. JACK) FULFORD in Mem. New York Bot. Gard. 11(4):
411. 1976.
Trail from Cochrane to Middleham Falls, secondary Vegetation, on rock in the middle of a small
road, rather exposed to sunlight, 600 m elev., c. per., 17997, det. VÄNA (PRC).
Distr.: Known from Brazil, Colombia, Mexico, Jamaica, Galapagos Islands, and recently reported
from Madeira by NlEUWKOOP & ARTS (1995, with fig.l:l-10). New to Dominica.
Ceratolejeunea flagelliformis (STEPH.) FULFORD in Bull. Torrey Bot. Club 71: 637. 1944.
Roseau Valley, elfin forest between Freshwater Lake and Boeri Lake, on branches of shrubs,
770 m alt., 17635; Morne Diablotin, elfin forest on western slope, epiphytic on shrub, 1230 m alt.,
17977.
Distr.: A relatively rare species, known from Puerto Rico, Martinique, Costa Rica, Dominica
(FULFORD 1987), and French Guiana (GRADSTEIN & HEKKING 1989).
 320

Ceratolejeunea patentissima (HAMPE & GOTTSCHE) A. EVANS in Bull. Torrey Bot. Club
32: 286. 1905.
Morne Diablotin, epiphytic in elfin forest on western slope, several collections between 1100 and
1260 m alt., 17968 (conf. GRADSTEIN), 17972, 17982; epiphytic on old tree in rain forest between
Cochrane and Middleham Falls, associated with Colura lyrata STEPH., 620 m, 18014/A.
Distr.: A small and easily oveiiooked species hitherto known from Puerto Rico, Cuba, Martinique,
Colombia (FULFORD 1945), Guyana, Surinam, French Guiana (GRADSTEIN & HEKKING 1989),
Ecuador (GRADSTEIN & al. 1994), and recently reported äs new to Dominica by SCHUSTER (1992)
where it seems to be not an uncommon species. All specimens are from well insolated sites.
*Cheilolejeunea adnata (KuNZE) GROLLE in J. Bryol. 9: 529. 1977.
= Cheilolejeunea decidua (SPRUCE) A. EVANS in Bull. Torrey Bot. Club 32: 188. 1905.
Roseau Valley, trau from Wotten Waven to Sulphur Springs, epiphytic near brook, 370-380 m,
c.spor., 18062.
Distr.: Widespread in the Neotropics (lowlands) from Brazil, Guyana, Surinam, French Guiana,
Venezuela, Colombia, Costa Rica, Mexico, Cuba, Puerto Rico, Bahamas, to Southern Florida.
Cheilolejeunea adnata was described and figured by EVANS (1906: pl. l, figs. 1-9, äs Cheilo-
lejeunea decidua (SPRUCE) A. EVANS), SCHUSTER (1980: figs. 688:13-14, 679), and LÜCKING
(1995: Abb. 27). The plants agree well with specimens collected in Brazil and determined by R.
GROLLE. Dominica lies well within the distribution area of this species.
* Colura cylindrica HERZOG in Svensk. Bot. Tidskr. 46: 106. 1952.
Rain forest near Emerald Pool, on rotten log, 400 m alt., 17730/A, conf. JOVET-AST (PC); Central
Forest Reserve above Beils, rain forest, epiphyllous on fern frond, 400 m alt., 17775.
Hitherto known from Ecuador (JOVET-AST 1953: 260), Brazil (SCHÄFER-VERWIMP 1989: 313),
Galapagos Islands (GRADSTEIN & WEBER 1982), Guyana, Surinam, French Guiana (GRADSTEIN
& HEKKING 1989), and recently reported from Tanzania, Africa, by Poes (1990); distribution
map in PÖCS (1991: fig. 9). New to Dominica and the West Indies.
Colura lyrata STEPH. in URBAN, Symb. Antill. 2: 471. 1901.
= Macrocolura lyrata (STEPH.) R. M. ScHUST. in J. Hattori Bot. Lab. 75: 233. 1994.
Morne Diablotin, elfin forest on western slope, epiphytic, 1250 m, 17973, conf. JovET-AST (PC);
rain forest between Cochrane and Middleham Falls, epiphytic on old tree, growing with Colura
rhynchophora, Harpalejeunea subacuta, Ceratolejeunea patentissima, and others, 650 m, 18002
(STU, Hb. ARTS), and 620 m, 18014 (EGR, G, JE).
This interesting species was hitherto known from Guadeloupe (type), Guyana (GRADSTEIN &
FLORSCHÜTZ-DE WAARD 1989; GRADSTEIN & HEKKING 1989), and Peru (SCHULTZE-MOTEL &
MENZEL 1987); recently reported from Dominica by SCHUSTER (1992) and EGGERS (1997, both
äs Macrocolura lyrata); obviously not a rare species on Dominica.
This species is kept here under Colura rather than under Macrocolura. SCHUSTER gives two
reasons for the generic Separation of Macrocolura one being the conspicuously botryoidal oil
bodies; however, already GRADSTEIN & al. (1977, with fig. 12) reported the presence of coarsly
segmented oil bodies in Colura patagonica AST (now a synonym of Colura nautnannii (SCHIFFN.)
STEPH., fide SOLARI 1983: 545), and, subsequently, very similar oil bodies of the Calypogeia type
were reported by LEON & al. (1998, with fig. 2: 2-3) for Colura omithocephala. An intermediate
condition between the wholly homogeneous, glistening oil bodies of „normal" Colura species and
the botryoidal oil bodies of Colura lyrata was found by SCHÄFER-VERWIMP (1992) from Colura
greig-smithii with finely but distinctly granulose-papillose oil bodies; these oberservations were
based surely on living plants. Therefore the different structure of the oil bodies cannot serve for
generic Separation of Macrocolura. In other genera of Lejeuneaceae, for example in
Aphanolejeunea, Lejeunea, Taxilejeunea, and Odontolejeunea, also occur homogeneous and
segmented oil bodies. The second reason for generic distinction fide SCHUSTER are the
symmetrical underleaves with erect Segments and a shallow, U-shaped sinus. In my opinion this
 321

character alone does not warrant generic Separation äs different types of underleaves also occur in
other genera, for example in Diplasiolejeunea. Theret'ore this species seems best kept in the
section Lingua of the genus Colura, äs placed by JOVET-AST (1953).
Colura rhynchophora AST in Rev. Bryol. Lichenol. 17: 27. 1948.
Morne Trois Pitons, on humid rotting log near brook, 550 m alt., 17755 (EGR, PC, STU); trail
from Cochrane to Middleham Falls, epiphytic in rain forest, c. per., 650 m elev., 18005/A (PC),
both conf. JOVET-AST.
Distr.: Obviously a rarely collected species, hitherto known only from Guadeloupe (type) and
Guyana (GRADSTEIN & FLORSCHÜTZ-DE WAARD 1989; GRADSTEIN & HEKKING 1989); very
recently reported from Dominica by EGGERS (1998).
Colura tenuicornis (A. EVANS) STEPH., Spec. Hep. 5: 942. 1916.
Roseau Valley, elfin forest between Freshwater Lake and Boeri Lake, epiphyllous on fern fronds,
c.per., 820 m, 17638 (PC), and epiphytic, c.per., 870 m alt., 17656/B (PC); Morne Diablotin, elfin
forest on western slope, epiphytic on shrubs, c.per., 1230 m alt., 17974/A (PC), all conf. JOVET-
AST.
Distr.: Nearly pantropical, distr. map in PÖCS (1991: fig. 8); surprisingly, EGGERS (1998) recorded
this locally abundant species on Dominica for the first time from the Island.
*Cyclolejeunea chitonia (TAYLOR ex LEHM.) A. EVANS in Bull. Torrey Bot. Club 31: 194.
1904.
Humid slope on road between Pont Casse and Emerald Pool, epiphytic on young Cecropia,
growing with Lejeunea flava (Sw.) NEES, and Drepanolejeunea lichenicola (SPRUCE) STEPH.,
500 m, 17721, det A. BERNECKER; Morne Diablotin, elfin forest on western slope, on dead branch
of shrub, 1080 m, 17950/B; epiphytic on Citrus in small Citrus plantation between Beils and Pont
Casse, 450 m, 17779.
Distr.: Costa Rica, Panama, Puerto Rico, Guadeloupe, Martinique, St. Vincent, Colombia,
Guyana, Suriname, French Guiana (LÜCKING 1995, with Abb. 33-34; figured also by ONRAEDT
1988 and BERNECKER-LÜCKING 1998b); known also from Brazil (HERZOG 1931, beside unpub-
lished records of the author).
The genus Cyclolejeunea, confined to the Neotropics in a number of about 20 species
(BERNECKER-LÜCKING 1998b: 176), is now known from Dominica with six species: C. accedens
(GOTISCHE) A. EVANS, C. chitonia (TAYLOR) A. EVANS, C. convexistipa (LEHM. & LlNDENB.)
A. EVANS, C. luteola (SPRUCE) GROLLE, C. mimula (STEPH.) STEPH., and C. peruviana (LEHM. &
LlNDENB.) A. EVANS. Of these, C. mimula is still not treated in a modern revision and may be
synonymous with another species. The genus is rieh on Dominica compared with five species
known from Costa Rica, a country famous for its diversity in hepatics (BERNECKER-LÜCKING
1998b).
*Cyrtolejeunea holostipa (SPRUCE) A. EVANS in Bull. Torrey Bot. Club 30: 553. 1903.
Rain forest on the track between Cochrane and Middleham Falls, epiphytic on old tree, 650 m,
c.per., 17999/A; Roseau Valley, trail between Wotten Waven and Sulphur Springs, epiphytic in
secondary Vegetation near brook, 370 m, 18059, det. GRADSTEIN (STU).
Distr.: Widespread in the Neotropics (Mexico, Honduras, Costa Rica, Colombia, Ecuador, Peru,
Venezuela, the Guianas, Brazil, Cuba, Puerto Rico, Guadeloupe, St. Vincent (LÜCKING 1995) but
hitherto not reported from Dominica.
Dactylolejeunea acanthifolia R. M. SCHUST. in Bull. Torrey Bot. Club 97: 343. 1970.
Rain forest on Emerald Pool, epiphyllous on fern fronds, c.per. and c.spor., (large population), ca
400 m alt., 17737 (BRNO, EGR, G, GOET, JE, KRAM-B, NY, PRC, STU, hb. ARTS; will be
distributed in the series „Bryophyta Neotropica Exsiccata", ed. by S. R. GRADSTEIN); trail from
Cochrane to Middleham Falls, in rain forest on fern fronds, ca 620 m alt., 18009 (JE).
 322

Hitherto known only from Dominica from where it was described and figured hy SCHUSTER
(1970). Locally frequent especially in the rain forest around Emerald Pool, occasionally elsevvhere
but easily overlooked. Always found on the fronds of a single fern species near the ground where
humidity is constantly high. GRADSTEIN (1992) considers this species äs endangered, and TAN &
al. (1994) in their World Red List of Bryophytes suspected it possibly äs extinct because „habitat
probably destroyed due lo hurricane"; however, äs our observations have shown Dactylolejeunea
acanthifolia may be more adequately placcd among the rare species. There is a good chance that
the rain forests on Dominica will not be destroyed by human activity, the single garantee for the
survival of this exciting hepatic. Hurricanes may not be a real danger for the survival of this
species because it is growing on small fern fronds in the undergrowth, and similar ecological
conditions may also occur in humid secondary Vegetation or in heavily damaged parts of old
growth rain forest.
Howcvcr, Phycolepidozia exigua R. M. SCHUST., considered, too, äs probably extinct by TAN &
al. (1994) could not be reJocatcd. Obviously, this species prefers the bark of old trees, and
hurricanes may be decisive for the survival of this monotypic genus.
•'Dicranolejeunea axiUaris (NEES & MONT.) SCHIFFN. in ENGLER & PRANTL, Nat.
Pflanzenfam. 1(3): 128. 1893.
Roseau Valley, trail between Freshwater Lake and Boeri Lake, hanging from shrub in elfin forest
along ridge, 810m, c.per., 17650.
Distr.: Common in Central America, in the tropical Andes and on the Galapagos Islands; rare in
the West Indies (Jamaica, Guadeloupe and Martinique); also on Mt. Roraima, Guyana (GRAD-
STEIN 1994). New to Dominica where it seems to be a rather rare species.
Diplasiolejeunea alata AST in Rev. Bryol. Lichenol. 17: 31. 1948.
Morne Trois Pitons, elfin forest on northwestern slope, epiphytic on small branches of shrubs,
c.per., 1030 m, 17829; Roseau Valley, on the track from Laudat to the Valley of Desolation,
humid ridge, epiphytic on shrub, c per., 880 m, 17875; Morne Diablotin, elfin forest on western
slope, epiphyllous on fern frond, c.per., 1230 m, 17975.
Distr.: Guadeloupe (type), Costa Rica, El Salvador, Brazil; very recently reported from Venezuela
by LEON & al. (1998); first and single record from Dominica by SCHUSTER (1970).
On Dominica Diplasiolejeunea alata is not a rare species and grows under quite similar ecological
conditions äs in Brazil where it was found often on small branches of shrubs (in secondary
Vegetation) at rather open and well insolated sites with high frequency of log (SCHÄFER-VERWIMP
1992).

Drepanolejeunea crucianella (TAYLOR) A. EVANS in Bull. Torrey Bot. Club 30: 33. 1903.
Humid secondary forest on roadside between Pont Casse and Emerald Pool, epiphyllous, 500 m
alt., 17723/D; rain forest on Emerald Pool, epiphyllous on fern fronds, growing between Cyclo-
lejeunea accedens (GOTTSCHE) A. EVANS, Aphanlolejeunea sicaefolia (GOTTSCHE) A. EVANS,
and Lejeunea flava (Sw.) NEES, 400 m alt., 17737/A, det. A. BERNECKER; Morne Trois Pitons,
rain forest on northwestern slope, epiphyllous, growing with Cyclolejeunea peruviana (LEHM. &
LINDENB.) A. EVANS and Aphanolejeunea cyathiphylla HERZOG, 1040 m alt., 17840/B, det. A.
BERNECKER (Hb. A. BERNECKER).
Distr.: Guyana, Surinam, Frcnch Guiana (GRADSTEIN & HEKKING 1989, GRADSTEIN & FLOR-
SCHÜTZ-DE WAARD 1989); recently reported by EGGERS (1997) from Dominica.
Drepanolejeunea lichenicola (SPRUCH) STEPH., Spec. Hep. 5: 335. 1913.
Road between Pont Casse and Emerald Pool, humid secondary Vegetation at roadside, a few plants
creeping on Cyclolejeunea chitonia (TAYLOR) A. EVANS, epiphytic on young Cecropia, 500 m
alt., 17721/A, det. A. BERNECKER.
Distr.: Mexico, El Salvador, Costa Rica, Panama, Colombia, Ecuador, Surinam, French Guiana,
Brazil, Jamaica, Puerto Rico, Guadeloupe (LÜCKING 1995, with fig. 45); Guyana (GRADSTEIN &
 323

FLORSCHÜTZ-DE WAARD 1989); firstly recorded for Dominica by SCHUSTER (1992, äs

„Drepanolejeiinea bispinulosa EVANS sensu amplo").
*Drepanolejeunea pinnatiloba SCHIFFN. in Bot. Jahrb. Syst. 23: 591. 1897.
Brook in rain forest on northwestern slope of Morne Trois Pitons, on very humid rotting log,
mixed with Colura rhynchophora, 550 m alt., 17754.
Distr.: Cuba (type), Puerto Rico (BISCHLER 1964), Guyana (GRADSTEIN & FLORSCHÜTZ-DE
WAARD 1989); BISCHLER (1964), äs well äs FULFORD (1987, dring BISCHLER 1964) excluded D.
pinnatiloba from the hepatic flora of Dominica äs the record by STEPHANI was based on two
misdetermined ELLIOTT collections. Therefore the above cited specimen which agrees perfectly
with the description and figures by BISCHLER (1964) is the first one from Dominica and the
Lesser Antilles.
Fossombronia bmsiliensis STEPH., Spec. Hep. 1: 382. 1900.
Roseau Valley, rain forest around Trafalgar Falls, on shady rock, 300 m alt., 17604; Roseau
Valley, between Wollen Waven and Sulphur Springs, on shady slope along road, 370 m all.,
18057, bolh wilh mature sporophyles.
Fossombronia brasiliensis is a ralher common species on roadside earlh banks in ihe Roseau
Valley Ihough there is only a single record by STEPHANI (1898-1900). Il is widespread in Ihe
Neotropics extending from Brazil norlhward lo Cenlral America, the West Indies (Puerto Rico,
Cuba, Haiti, Dominican Republic, Trinidad) and to Ihe soulheastern United Slales (SCHUSTER
1992, wilh figs. 843:16-18, 844:1-6, 845).
Haplomitrium blurnii (NEES) R. M. SCHUST. in J. Hattori Bot. Lab. 26: 225. 1963.
= Haplomitrium andinum (SPRUCH) R. M. SCHUST. in J. Hattori Bot. Lab. 26: 225. 1963
Roseau Valley, track from Laudat to Middleham Falls, rain foresl near Ihe falls, on shady rock
over a Ihin layer of humus, in large quanlity, 620 m alt., 17703 (EGR, GE, STU, hb. ARTS);
Morne Trois Pitons, rain forest on northweslern slope, on humid rolling wood, 860 m alt., 17834;
Morne Diablotin, rain foresl on weslern slope, on decayed log, 800 m elev., 17926.
Dislr.: Known äs H. andinum from Ecuador, Peru, Guadeloupe, Martinique, Dominica; under Ihis
name lisled äs a „rare species" by GRADSTEIN (1992); after SCHUSTER (1966a: 56) „apparantly a
rather rare species" and found by him only in a „Irace" on Dominica from where il is cited in the
literalure only from Iwo old colleclions; äs H. blumii known from Iropical Asia and recently also
reported from topical Africa (GROLLE 1993, with figs. 1+2). The three colleclions cited above
show that this species is surely not a rare one on Dominica - the population near Middleham Falls
reachcd aboul an area of 30 x 40 cm in an almost pure mal - and lhal il may be found on Ihis
island elsewhere.
*Harpalejeunea emarginata AST in Rev. Bryol. Lichenol. 16: 38. 1947.
Rain foresl on Irail from Cochrane lo Middleham Falls, epiphylic on large iree, growing wilh a
few planls of Anoplolejeunea conferta (MEISSN.) A. EVANS, Drepanolejeiinea crassiretis A.
EVANS, and Cheilolejeunea rigidula; 650 m, 17999/B, del. GROLLE (JE).
Dislr.: Hilherlo known only from Ihe type collection from Guadeloupe (JOVET-AST 1947, wilh fig.
IX.). New lo Dominica.
*Harpalejeunea subacuta EVANS in Bull. Torrey Bot. Club 30: 547. 1903.
Rain foresl on Ihe track from Cochrane to Middleham Falls, epiphytic on large Iree, growing wilh
Colura lyrata and Colura rhynchophora; 650 m, 18005/B, del. GROLLE (JE).
Dislr.: Puerto Rico (EVANS 1903, with figs. 1-11 on pl. 20) and El Salvador (WINKLER 1967);
firsl record for Dominica and Ihe Lesser Anlilles.
*Lejeunea filipes SPRUCE in Trans. & Proc. Bot. Soc. Edinburgh 15: 275. 1884. [„£«-
Lejeunea"}
= Lejeunea autoica R. M. SCHUST. in J. Hattori Bot. Lab. 25:6. 1962, syn. fiele REINRR-DREHWALD 1995: 127.
 324

Roseau Valley, rain forest on trail from Laudat to Middleham Falls, epiphytic near brook, 550 m
alt., 17687 p.p.; secondary Vegetation above Giraudel (south of Roseau), epiphyllous, 650 m alt.,
17912, both det. REINER-DREHWALD.
Distr.: USA (Florida), Cuba, Peru (type), Argentina, Cameroon, Ghana, and Tanzania (REINER-
DREHWALD 1995); Colombia (WINKLER 1976: 797, 814), and Costa Rica (DAUPHIN & al. 1998,
äs L. autoica); distribution map in GRADSTEIN & al. (1983, äs Lejeunea autoica). The species was
described and figured by JONES (1979: fig. 2, äs L. autoica), SCHUSTER (1980: figs. 716-717, 719:
15-17, äs L. autoica), and REINER-DREHWALD (1995: fig. 23, 53C); Lejeunea filipes commonly
grows epiphytic, more rarely on living leaves.
*Lejeunea (Crossotolejeunea) paucidentata (STEPH.) GROLLE in J. Hattori Bot. Lab. 69:
191. 1991.
= Crossotolejeunea paucidentata STEPH. in Hedwigia 35: 76. 1896.
Above Roseau, on the track from Cochrane to Middleham Falls, rain forest, on rotting fern stem,
620 m alt., 17998, det. A. GODA.
Distr.: Hitherto known only from Cuba (GROLLE 1991). New to Dominica and the Lesser Antilles.
*Lejeunea tapajosensis SPRUCE in Trans. & Proc. Bot. Soc. Edinburgh 15: 223. 1884.
[„Taxi-Lejeunea"]
Secondary Vegetation above Giraudel (south of Roseau), epiphytic on young tree in pasture,
520 m alt., 17900/A, det. REINER-DREHWALD.
Distr.: Brazil (including type), Peru (SCHULTZE-MOTEL & MENZEL 1987), Argentina (REINER-
DREHWALD 1995). Seems to be the first record for the West Indies and the first one outside
Continental South America. The species was described and figured by REINER-DREHWALD (1995:
fig. 35).
*Leptolejeunea elliptica (LEHM. & LiNDENB.) SCHIFEN. in ENGLER & PRANTL, Nat.
Pflanzenfam. 1(3): 126. 1893.
Central Forest Reserve near above Beils, rain forest, epiphyllous, growing with Drepanolejeunea
inchoata (MEISSN.) SCHIFFN., 400 m alt., 17777, det. A. BERNECKER [„form with subacute leaf'].
Distr.: Widespread in the Neotropics but hitherto not reported from Dominica.
Leptolejeunea radicosa (NEES ex MONT.) GROLLE in J. Hattori Bot. Lab. 45: 178. 1979.
= Leptolejeunea obovata BlSCHL. in Nova Hedwigia J7: 319. 1969.
Humid slope on roadside between Pont Casse and Emerald Pool, epiphyllous on fern fronds,
500 m alt., 17723/C, det. PÖCS (äs L. obovata); rain forest around Emerald Pool, epiphyllous,
c.per., 400 m, 17727/C; rain forest on the track from Cochrane to Middleham Falls, epiphyllous
on fern fronds, growing with Dactylolejeunea acanthifolia, Aphanolejeunea sicaefolia, Cyclo-
lejeunea convexistipa, 620 m, 18011.
Distr.: Jamaica, Panama, Colombia, French Guiana (BtSCHLER 1969, äs L. obovata, with fig. XI
and distr. map Carle IV), Costa Rica (MORALES 1991, äs L. obovata). Recently reported from
Dominica by EGGERS (1997).
The genera Leptolejeunea and Cololejeunea are surprisingly rare on Dominica. We have collected
only a few specimens of both these genera whereas different species of other genera of
Lejcuneaceae, for example Diplasiolejeunea, Cyclolejeunea, Drepanolejeunea, Prionolejeunea,
Colura, and Aphanolejeunea, have been seen very frequently.
*Microlejeuena acutifolia STEPH. in Hedwigia 35: 113. 1896.
Above Roseau, on the track from Cochrane to Middleham Falls, rain forest, epiphytic on small,
rotting stem, growing with Cheilolejeunea rigidula (NEES ex MONT.) R. M. SCHUST., Lejeunea
laetevirens NEES & MONT., and Echinocolea sp.; 650 m alt., 18017/A.
 325

Distr.: Mexico, El Salvador, Costa Rica, Cocos Island, Guianas, Colombia, Puerto Rico,
Guadeloupe, Martinique, St. Vincent, Trinidad (LÜCKING 1995, with Abb. 57A-G, äs Lejeunea
(Microlejeiinea) acutifolia', first record for Dominica.
'''•Plagiochila steyermarkii H. ROB. in Bryologist 68: 93. 1965.
Morne Diablotin, elfin forest on western slope, epiphytic on shrub, 1270 m, 17964, det.
GRADSTEIN (JE).
Distr.: This remarkable species was hitherto known from the type specimen from Auyan-tepui
from Venezuela (ROBINSON 1965) and Guyana (GRADSTEIN & FLORSCHÜTZ-DE WAARD 1989).
INOUE (in GRADSTEIN & FLORSCHÜTZ-DE WAARD 1989) reduced the species to a synonym of
Plagiochila aerea TAYLOR; however, according to GROLLE & HEINRICHS (1999) this is erro-
neous. New to the West Indies.
*Pycnolejeunea contigua (NEES) GROLLE in J. Hattori Bot. Lab. 45: 179. 1979.
Rain forest on the track from Cochrane to Middleham Falls, epiphytic, 620 m alt., 18015 (c.per.),
18008, conf. X.-L. HE.
Distr.: Brazil, Costa Rica, Peru, Ghana, Madagaskar (GROLLE 1979), Trinidad; Sierra Leone,
Western Nigeria,Tanzania (JONES 1979); distr. map in GRADSTEIN & al. (1983: pl. LI, fig. 7). The
species was described and figured by HERZOG (1951, äs P. densistipula var. dubia), KACHROO &
SCHUSTER (1961, äs P. spruceana and P. papulosa), and JONES (1979: fig. 6).
Both specimens of Pycnolejeunea contigua from Dominica were collected from the stem of large
trees (1-2 meters above ground) along tracks in undisturbed primary rain forest. In Brazil this
species has been found by the author and bis wife in the states of Santa Catarina, Säo Paulo and
Minas Gerais from sea level to 1440 m, in a variety of different woodlands, in mangrove- and
restinga-vegetation äs well äs in cacao-plantations, in undisturbed rain forests and even in semi-
decidous mountain forests. The collection sites were often well insolated though regularly in the
undergrowth, and obviously P. contigua prefers humid ecological conditions. JONES (1979) found
P. contigua in West Africa only on the branches in the crowns of large trees in rain forest, and not
at all in the drier mixcd deciduous forest.
*Radula mazarunensis YAMADA in Trop. Bryol. 1: 38. 1989.
Central Forest Reserve above Beils, on humid earth slope along road, c.per., 400 m alt., 17767,
det. YAMADA (hb. YAMADA).
Distr.: Hitherto known only from the type collection from Guyana (YAMADA 1989). New to the
West Indies and the first record outside Continental South America.
*Radula subinflata LiNDENB. & GoTTSCHE in GoiTSCHE, LINDENBERG & NEES, Syn.
Hep.: 724. 1847.
Roseau Valley, trail from Laudat to Middleham Falls, submerged on dripping rocks near the falls,
c.per., 620 m alt., 17707, det. YAMADA (hb. YAMADA); Morne Trois Pitons, epiphytic in rain
forest on northwestern slope, 700 m elev., 17787.
Distr.: Widely distributed in the Neotropics including Mexico, Guatemala, West Indies (Guade-
loupe, Martinique), Surinam, Venezuela, Ecuador, Galapagos Islands, Brazil (YAMADA &
GRADSTEIN 1991), and Colombia (URIBE & GRADSTEIN 1998). First record for Dominica.
*Radula tenuis YAMADA in J. Hattori Bot. Lab. 54: 247 1983. Fig. l
Rain forest on Freshwater Lake, epiphytic, 750 m, 18048 (c.per.) and 18051, both det. YAMADA
(hb. YAMADA).
Distr.: Hitherto known only from the type collection from Cuba (YAMADA 1983: 247, 1988: 383).
The specimen 18048 seems to be the first one with perianths: Perianth mostly 2 mm long,
trumpet-shaped in outline, the base ca. 0,2 mm wide, terete, above flattened and broadened, the
mouth ca. 0,75 mm wide, faintly sinuate-undulate; the cells in the central and basal part
±rectangular, 20-35 u.m long, 12-20 um broad, all cells with one large oil body, cells at mouth
smaller, subquadrate, 10-15 x 10-18 um; mature sporophyte not seen (fig. 1).
 326

Fig. l : Radula tenuis YAMADA A: portion of plant with perianth, ventral view; scale = 0,5 mm.
B: perianth with female bracts, dorsal view; scale äs A. C: upper portion of perianth showing
faintly sinuate-undulate mouth; scale = 0,25 mm. D: cells of perianth mouth; scale = 60 um.
E: median cells of perianth, with oil bodies; scale = 10 um.

*Syzygiella integerrima STEPH., Spec. Hep. 6: 117. 1917.

Roseau Valley, trail from Laudat to Valley of Desolation, epiphytic in humid shrubby Vegetation
along ridge, 840 m alt., c.per., 17872.
Distr.: Brazil, Guadeloupe (INOUE 1966), and Colombia (GRADSTEIN & al. 1977). New to
Dominica.

Musci

Brymela websteri (CRUM & BARTR.) BÜCK in Brittonia 39: 218. 1987.
Roseau Valley, trail from Laudat to Valley of Desolation, in rain forest on moisl wood of a small
bridge, 700 m, 17861, det. BÜCK (NY, STU).
Distr.: Brymela websteri is known only from Panama, Jamaica, and Dominica; the record from
Dominica in BÜCK (1998, with figs. 1-8 on pl. 20) is based on the single specimen cited above.
 327

*Biyum Umbatum MÜLL. Hai, Syn. Muse. Frond. 2: 573. 1851.

Roseau Valley, trail from Laudat to Middleham Falls, on rock in brook, sometimes submerged,
500 m alt., 17677; Roseau Valley, above Wollen Waven and Sulphur Springs, on rock in brook,
sometimes submerged, 370 m alt., 18068.
Widespread in the Neolropics including Mexico, Costa Rica, Cuba, Haili, Dominican Republic,
Grenada, Guadeloupe, Trinidad, Colombia, Venezuela, Ecuador, Peru, Bolivia, and Brazil (OCHI
1980, with fig. 36A-J; figured also in SHARP & al. 1994: fig. 347a-g). New lo Dominica.
*Calvmperes venezuelanum (MlTT.) BROTH. ex PlTTlER in Bol. Soc. Venez. Ci. Nat. 3:
360. 1936.
Morne Trois Pitons, epiphytic in rain forest on western slope, 900 m alt., 17832.
Distr.: Panama, Jamaica, St. Vincent, Guadeloupe, Venezuela, Trinidad, Guyana, Suriname,
French Guiana (REESE 1993, with distribulion map and figs.). New to Dominica.
*Daltonia longifolia TAYLOR in London J. Bot. 7: 284. 1848.
Roseau Valley, on the track from Laudat to Freshwater Lake, culture zone, epiphytic on Citrus,
associated with Daltonia steiwpltvlla MlTT. and Actinodontium integrifolium (BROTH.)
CHURCHILL, 700 m, 17628/B; Morne Diablotin, clfin forest on western slope, epiphytic on rotting
stem, growing with Daltonia stenophylla, Trachyxiphium guadalupense (BRID.) BÜCK, Leskeodon
auratus (MÜLL. Hai.) BROTH. and some small hepatics, 1250 m alt., 17970/A, both collections
only in small quantity.
Distr.: Mexico to Panama, Venezuela and Colombia to Bolivia, Cuba, Jamaica, Haiti, Dominican
Republic, Puerto Rico, Guadeloupe, Martinique; first record for Dominica.
Daltonia stenophylla Mm. in J. Linn. Soc., Bot. 12: 402. 1869.
Roseau Valley, on the track from Laudat to Freshwater Lake, culture zone, epiphytic on Citrus',
growing with Daltonia longifolia and Actinodontium integrifolium, 700 m alt., 17628/A; ibid,
track from Laudat to Valley of Desolation, epiphytic in humid shrubby Vegetation, 880 m elev.,
17878; Morne Diablotin, elfin forest on western slope, epiphytic on rotting stem, admixt with
Trachyxiphium guadalupense and Leskeodon auratus in a collection of Daltonia longifolia, 1250
m alt., 17970/A p.p. — all three specimens in small quantity.
Distr.: Costa Rica to Boliva, Brazil, Cuba, Jamaica, Dominican Republic, Puerto Rico, St.
Vincent; seems to be only twice collected on Dominica (see TOWNSEND 1994). An easily over-
looked and therefore undercollected species, obviously not seen from Dominica by BÜCK (1998,
with figs. 1-6 on p]. 1).
Entosthodon bonplandü (HOOK.) MlTT. in J. Linn. Soc., Bot. 12: 245. 1869.
Morne Diablotin, elfin forest on western slope, on humid earth bank, c. spor., associated with
Alobiella husnotii (GOTTSCHE) SCHIFFN., 1070 m, 17947.
Distr.: Venezuela, Colombia, Ecuador, Galäpagos, Peru, Bolivia, Paraguay, Brazil, Mexico,
Guatemala, Costa Rica, Jamaica, Dominican Republic, Puerto Rico, Martinique, Trinidad, (FlFE
1987, with figs. 50-70). Entosthodon bonplandü was seen several times on Dominica but only
once collected; it seems not to be quite rare though reported firstly for the island from two
gatherings by TOWNSEND (1994).
*Epipterygium wrightii (SÜLL.) LlNDB. in Öfvers. Förh. Kongl. Svenska Vetensk.-Handl.
19: 604. 1863.
Roseau Valley, rain forest near Trafalgar Falls, on shady rocks, 280 m, 17617.
Distr.: West Indies (Cuba, Jamaica, Dominican Republic, Haiti, Puerto Rico, Martinique),
Venezuela, and Peru. After SHAW (1984, with key and figures) this species is common in
the Greater Antilles but has been found only once in the Lesser Antilles. New to Dominica
being the second record for the Lesser Antilles.
 328

*Leucobryum albicans (SCHWAEGR.) LlNDB. in Öfvers. Förh. Kongl. Svenska Vetensk.-

Handl. 20: 402. 1863.
Morne Diablotin, elfin forest on western slope, on shady rock, 1260 m alt., 17967.
The plants have leaves more or less 12 mm long and the leucocysts at leaf base are only in two
layers with rarely a few cells in a third layer - exactly äs figured by YANO (1975: fig. 18h) and
described by COSTA (1989); however, YANO described the leaf bases äs three- to four-layered and
figured it two-layered, and COSTA (1989) described them äs two-layered, sometimes three-
layered, and figured it three- to four-layered (COSTA 1989: fig. 3e).
Distr.: West Indies (Cuba, Puerto Rico), Central- and South America (fide Index Muscorum also
As l). First record for Dominica and the Lesser Antilles.
*Sphagnumperichaetiale HAMPE in Linnaea 20: 66. 1847.
Morne Trois Pitons, elfin forest on western slope, on humid rock, 1040 m alt., 17827 (MICH);
Roseau Valley, trail from Laudat to the Valley of Desolation, on ridge, at several sites between
750 and 950 m alt., 17869, both det. CRUM (G, MICH, STU).
Distr.: S. perichaetiale is widespread in the Old and New World tropics and subtropics (distr. map
in EDDY 1977) under many names (EDDY 1977 listed 52 synonyms). From the West Indies it is
known from Cuba, Jamaica, Puerto Rico, Guadeloupe, and Trinidad. The genus Sphagnum is new
to Dominica.

Discussion

The number of bryophytes known from Dominica is near 430, about 270 taxa of hepatics
in 85 genera and 27 families, and 160 taxa of mosses in 78 genera and 25 families
(SCHÄFER-VERWIMP, msc.). In this paper 32 species are newly reported, 26 hepatics and 6
mosses (including each one new family of hepatics and mosses, Cephaloziellaceae and
Sphagnaceae, four new genera of hepatics, Cephaloziella, Cyrtolejeunea, Dicrano-
lejeunea, Pycnolejeunea, and two new genera of mosses, Epipterygium and Sphagnum),
leading the total number to approximately 460. The bryophyte flora of Dominica is
relatively well known due to the extensive collections made by ELLIOTT more than
hundred years ago, but still far from complete, and the expected number of species may
well lie above 500.
Seven of the presently reported species are new to the West Indies: Aphanolejeunea
cingens, A. costaricensis, A. cyathiphylla, Colura cylindrica, Radula mazarunensis,
Lejeunea tapajosensis, and Plagiochila steyermarkii, for the latter two species this is the
first record outside Continental South America. New to the Lesser Antilles are Aphano-
lejeunea camillii, Cephaloziella granatensis, Cheilolejeunea adnata, Drepanolejeunea
pinnatiloba, Harpalejeunea subacuta, Lejeunea filipes, Lejeunea paucidentata, Pycno-
lejeunea contigua, Radula tenuis, Brymela websteri, and Leucobryum albicans. Some
species may have been expected to occur also on Dominica äs they are known from
neighbouring Islands; to these belong Aphanolejeunea verrucosa, Cephaloziella antillana,
Cyclolejeunea chitonia, Cyrtolejeunea holostipa, Microlejeunea acutifolia, Radula subin-
flata, Bryum limbatum, and Sphagnum perichaetiale. Rare species (at least on Dominica)
belonging to this category are Dicranolejeunea axillaris, Harpalejeunea emarginata,
Leptolejeunea elliptica, Syzygiella integerrima, Calymperes venezuelanum, Daltonia
longifolia, and Epipyterygium wrightii. Finally, some rarely collected or easily overlooked
species could be confirmed to occur on the island: Acanthocoleus aberrans var. aberrans,
Aphanolejeunea sicaefolia, Arachniopsis diacantha, Ceratolejeunea flagelliformis,
 329

C. patentissima, Colura lyrata, Dactylolejeunea acanthifolia, Diplasiolejeunea alata,

Drepanolejeunea lichenicola, Fossombronia brasiliensis, Haplomitrium blumii, Daltonia
stenophylla, and Entosthodon bonplandii. Further five species which were reported very
recently during compiling this list could be confirmed, too: Aphanolejeunea angustissima,
Colura rhynchophora, C. tenuicornis, Drepanolejeunea crucianella, and Leptolejeunea
radicosa.

Acknowledgements
For Identification or confirmation of specimens I very like to thank Drs A. BERNECKER-LÜCKING, W. R.
BÜCK, H. CRUM, A. GODA, S. R. GRADSTEIN, R. GROLLE, X.-L. HE, S. JOVET-AST, E. REINER-DREHWALD,
J. VÄNA, and K. YAMADA. S. R. GRADSTEIN is gratefully acknowledged for reviewing the manuscript, and P.
GEISSLER and W. R. BÜCK for providing literature and for the loan of specimens from G and NY.

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Author: Alfons SCHÄFER-VERWIMP, Mittlere Letten 11, D-88634 Herdwangen-Schönach

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