Sérgio, Cecilia

Revistes Catalanes amb Accés Obert

Orsis, 3: 5-13 (1988) 
Morphological, karyological and phytogeographjc 
observations on Entosthodon curvisetus (Schwager.) 
C .  Müll. as a basis for a new genus, Funariella Sérgio 
(Funariaceae: Musci) 
Cecilia Sérgio 
Museu, Laboratório e Jardim Botlnico. Faculdade de Ci&ncias de Lisboa. 
Rua da Escola Politécnica. 1924 Lisboa Codex (Portugal). 
Key words: Entosthodon curvisetus , ecotogy, karyology, morphology, phytogeography, scan- 
ning photographs spores, Funariella gen. nov. 
Abstract. Based on comparative studies o£ sporophyte characters such us capsule morphology, 
seta length, arrangement of cells in the operculum, absence of peristome, spore morphology 
and chromosome number (n=5) a new genus has been proposed, Funariella Sérgio for the 
Funariaceae. The european material of F. curviseta has been revised and the ecology and 
general distribution are discussed briefly. 
Resumen. Observaciones morfológicas, cariológicas y fitogeográficas sobre Entosthodon curvi- 
setus (Schwaegr.) C. Miill. como base para la descripción de un género nuevo Funariella Sérgio 
(Funariaceae: Musci). Se propone un nuevo género, Funariella Sérgio, en base al estudio 
comparativo de 10s caracteres del esporófito tales como la morfologia de la cápsula, longitud 
de la seta, disposición de las células del opérculo, ausencia de peristoma, morfologia de las 
esporas y número cromosómico. 
Introduction 
In connection with the revision of some Mediterranean species of bryophytes 
for preparing distribution maps to be included in aworkgroup for mapping 
the bryophytes in Europe,,, approximately 50 specimens of Entosthodon 
curvisetus (Shwaegr.) C. Müll, from the following herbaria (LISU, LISFA, 
INA, G, BM, PC, BCB, MA, FI, DUIS, HEBRARD) have been exami- 
ned. Additionally, as a result of severa1 bryological explorations in calca- 
reous areas of Portugal (Sérgio et. al. 1984, Sérgio & Sim - Sim 1985), many 
samples of this interesting Funariaceae, always considered a rare species, 
were collected. For these reasons, a short revision of this taxon has been 
undertaken. 
Generic concept 
A generic revision of the Funariaceae was recently published (Fife 1985) 
based especially on numerical phenetics studies, spore morphology and 
evolutionary trends. In this work E. curvisetus was provisionally placed in 
Entosthofdon section Entosthodon with some reservations, althougt this 
taxon presents phenetic similarities in sporophytic characters nearer the 
subgenus Plagiodus Fife. 
The majority of the distinctive characters of the Funariaceae are in 
sporphytes (Fife 1985) and the morphological features of E. curvisetw show 
that this taxon had some intermediate characteristics between the subg. 
Entosthodon and the subg. Plagiodw, and presents a similarity coeficient 
considered by Fife as a monotypic genera or subgenera. 
Figure 1. Spores of Funariella curviseta (Schwaegr.) Skrgio wmb. nov. Scanning electron 
micrographs. Specimens 1.2 Algeria, 1904, Dixon (BM); 3.4 Italy, Sardinia, 1832, Muller 
(BM). Scale bar = 10 pm. 
I have been studing this taxon since 1989, particulary the spore ornamen- 
tation, that shows considerable differences from other Funariaceae by ha- 
ving a distinct reticulum whit minutely verrucate-baculate perine processes 
that border more or less regular alveoli (Fig. 1.). Limpricht (1885-1903) first 
mentioned the presence of <<dicht mit klein Netzfaltenn in these spores, and 
Roth (1904) also mentioned ccnetzfalting oder locker warzig und durchs- 
cheinend,, . 
Recently Fife (1985), studing 53 taxa if the Funariaceae with electron 
Figure 2. Funariella curviseta. (Schwaegr.) Sérgio comb. nov. 1 Leaf; 2 Leaf median margin; 3 
Leaf apice; 4 Axillary hair; 5 Immature sporophyte; 6-7 Calyptra; 8 Mature capsule; 9 Paraphy- 
ses and antheridia; 10-11 Arrangement of operculum cells; 12 Dry plant with empty capsule; 
13 Longitudinal cross section of capsule mouth; 14 Capsule mouth; 15 Exothecial cells; 16 
Stoma. Specimens: 1-16 Portugal: Serra da Arrábida, Alpertuche, C. Sérgio & M. Sim-Sim 
4769 (LISU); 11 Corsica: Cap. Corse, J.P. Hébrard; 916.4.76 (LISU). 
microscopy, showed that E. curvisetus spores presents an ornamentation 
that occurs in no other species of this genus. 
In addiction there is no infraspecific variation in this taxon and the 
sporophyte characters are stable. Also, E. curvisetus is easily distinguished 
from others species of the genus by the seta length (1.2 to 2 mm), its ratio 
with the capsule (from 0.8:l to 1:0.8), the asymmetric, inclined to pendent 
capsule before dehiscence, the absence of peristome and the marked spiral 
arrangement of cells in the operculum (Fig. 2). 
The spiral arrangement of the operculum cells occurs only in Entosho- 
don in the subg. Plagiodus sensu Fife and this Mediterranean species has an 
operculum with this character (Fig. 2). Schimper (1856) has described the 
operculum of E. curvisetus as having series cellularum oblique versus cen- 
trum convergentes. On the other hand the species of this subg. Plagiodus 
have the capsule with double peristome or a well developed exostome, 
while E. curvisetus has no peristome. The operculum on the species of the 
subg. Entosthodon presents irregularly arranged cells. The capsules are 
erect and more or less symmetric. 
In Bryologia Europaea, Bruch et al. (1836-1851) referred to an other 
interesting characteristic of the sporophyte. The calyptra when young is 
fusiformis tetragona, related to the genus Pyramidula or Gonimitrium that 
have a persistent and distinctly 4-8 angled calyptra. However in E. curvise- 
hrs this feature is observed only in the first phase of development of the 
sporophyte and is not a constant calyptra characters of E. curvisetus in 
herbarium specimens. 
Karyology 
As the sporophyte morphology suggests a possible segregation of E. curvi- 
setus from others species of this genus, chromosome studies were caried out 
on this taxon in 1983 and 1984, with fresh material from Portugal to try to 
confirm these differences. 
Chromosomes were counted at meiosis ahd the number of bivalents 
determined. Only 5 could be discerned, with apparent size ca. 2.5-4pm 
(Fig. 3). 
Chromosome studies have been made on all classes of bryophytes but 
the number of species has been small (about 3 % to 14 % in different 
. groups, after Smith 1978). However, a detailed treatment of bryophyte 
karyology was made by Fritsch (1982) and studies of cytogenetics and evo- 
lution in the more important groups, genera and also species, have been 
published recently (Iwatsuki & Inoue 1984, MC Adam 1982, Ramsay 1983a, 
Smith 1978, Steere 1972). 
A remarkable diversity in chromosome numbers in moss species was 
found with the lowest value n=4 (Ramsay 1983b). Species with n=5 are 
rare with some examples in Fissidens (Iwatsuki & Inoue 1984) while high 
Figure 3. Photomicrographs of sporocystes of Funariella curviseta at meiosis (acetato-orcein 
stein, with Geimsa's 6-8 pH method). Bivalents 5 -+n=5. Specimen Portugal, Serra da Arrá- 
bida, Alpertuche, 2.03.1983, C. SCrgio & M. Sim-Sim 4769 (LISU). 
numbers of n =10 to n=13 are more frequent in the Bryopsida (Smith 1978). 
A large part of the Funariaceae species still remain totally unknown in 
terms of their chromosome numbers (Fife 1985), and the tropical and me- 
diterranean taxa have not yet been studied. However, Steere (1972) consi- 
dered the basic numbers n=9, n=12, n=13 or n=14 for this family, while 
Smith (1978) refered to the haploid number of the Funariaceae as n=7 or 
n=9. This author suggests that the elements of this family evolved from an 
earlier stock than other Diplolepideae, before the chromosome number 
of 6 had been derived. 
~ 
A species of Nanomitrium, an element of a family related to the Funa- 
riaceae, has a haploid chromosome number n=10, being the basic number 
of a large series of Bryaceae (Steere 1972) and Fissidentaceae (Iwatsuki & 
Inoue 1984). 
Based on the acquisition of this detail of E. curvisetus I think that this 
species has a different karyological base to segregate it from the rest of the 
Entosthodon genus, that has n=14, n=9 or its multiples (Fife 1985). 
It is possible that in this taxon karyological changes were involved in its 
evolution, which agrees with the cytological phylogenies suggested by Smith 
in the Diplolepideae-Acrocarpae group (n=6 -, n=5 -, n=10). Then spe- 
cies with n=5 where derived from the ancestral stock which has n=6. 
On the other hand, with the principles of moss systematics of Miller 
(1979) it is concluded that the sporophyte of E. curvisetus may involve an 
evolutionary advance with short seta and peristome absent. The monoecious 
condition of this moss also represents phylogenetic evolution. 
Funariella gen. nov. 
Capsula subpendula longicolla subgibbosa in pedicello breviore arcuat0 ho- 
rizontalis; sicca vacua macrostoma. Operculum plano-convexum. Operculi 
series cellularum oblique versus centrum convergentes. Peristomium nullum. 
Sporae ferruginae 20-24 pm diam., manifeste reticulata, muri verrucoso-ba- 
culatae. Chromosomatum numerus n=5. 
Type species: Funariella curviseta (Schwaegr.) Sérgio comb. nov. 
Basionym: Gymnostomum curvisetum Schwaegr., Spec. Musc, Suppl. 2 
(1): 17, Tab. 105 (1823). 
Main synonymy: Physcomitrium curvisetum Brid., Bryol. Univ. 11: 815 
(1827). Entosthodon curvisetus (Schwaegr.) C. Muller, Syn. 1:121 (1848). 
Funaria curviseta Milde, Bryol. siles. : 196 (1869). 
Illustrations: Schwaegrichen. 1823. Tab. 105. Bruch, P., W. Ph. Schim- 
per & T. Gümbell. 1836-1855. Tab. 301. Husnot, T. 1884-1890. Tab. 58. 
Roth, G. 1904. Tab. 44.2. Brotherus, V. F. 1924-1925.: 329 fig. 278. 
Ecology and habitat 
Funariella curviseta has a termophytic character and is found mostly on wet 
sites with maritime influence. It prefers saxicolous substrata in fissures of 
rocks and grows frequently on calcareous areas. 
In Portugal, this species is usually found mixed with Mediterranean 
elements, sometimes with rare species such as Fissidens intralimbatus (Sér- 
gio & Sim-Sim 1985) and in the same communities reported by Hébrard 
(1984) from Corsica. 
The great majority of the European records of this Funariaceae are in 
areas whit mean annual temperatures 15" C or more. 
Distribution 
The Funariaceae, which is a world-wide family consisting of about 276 
species in 13 genera, has the great majority of its taxa restricted to the 
northern temperate zones, althougth there is a great diversification in tropi- 
cal Africa with 11 genera (Ochyra 1983), while Europe has only 6 genera. 
F. curviseta has a Mediterranean-macaronesian distribution pattern. 
Most european records are in areas near the coast or in islands (Fig. 4). It 
is distributed from the eastern part of the Mediterranean basin (Syria, 
Crete, Eggypt and Cyprus) to the Iberian Peninsula and Canary Islands 

(During 1981, Eggers 1982). Nevertheless, it always a restricted occurence, 
and more than 40 % of all its collections were made in the last century. 
In the Mediterranean region, many monotypic bryophyte genera are 
present and endemic species are included in some Funariaceae such as in 
Gonimitrium and Pyramidula that are also a strongly isolated group in this 
family, and exhibit many morphological features which can be considered 
allied to the Gigaspermaceae (Fife 1982) which also has endemic Medi- 
terranean taxa. 
The isolation of F. curviseta from its congeners confirms its recognition 
as a monotypic genus. 
Acknowledgements 
I wish to thank the director of E.M. Unit of British Museum who gave me the opportunity of 
prepar the scanning micrographs of spores and P.M. Jones of King's College for his reading of 
the manuscript. 
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Manuscript received on November 1986. 


Morphological, karyological and phytogeographic observations on Entosthodon curvisetus (Schwager.)C. Müll. as a basis for a new genus, Funariella Sérgio (Funariaceae: Musci)

Morphological, karyological and phytogeographic observations on Entosthodon curvisetus (Schwager.)C. Müll. as a basis for a new genus, Funariella Sérgio (Funariaceae: Musci)

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