Contribution to the biosystematic study of liverworts (Marchantiophyta) of Senegal:
Case of Senegaleses herbaria and check-lists
Djibril Diop 1, *, Mamadou Sydibé 1, Ali Mohamed Abdou Salam 1, Doudou Diop 2, Seydina Diop 2, Jules Diouf 1,
Abdoul Aziz Camara 1, Mame Samba Mbaye 1 and Kandioura Noba 1
Botany - Biodiversity Laboratory, Department of Plant Biology, Faculty of Science and Technology, Cheikh Anta Diop
University, BP 5005, Dakar-Fann, Senegal.
2 Fundamental Institute of Black Africa Cheikh Anta Diop (IFAN-CAD) Dakar- Senegal.
1
GSC Biological and Pharmaceutical Sciences, 2021, 17(03), 078–084
Publication history: Received on 30 October 2021; revised on 10 December 2021; accepted on 12 December 2021
Article DOI: https://doi.org/10.30574/gscbps.2021.17.3.0342
Abstract
The aim of this work is to contribute to a better knowledge of liverworts in the herbaria of Senegal in order to provide
basic knowledge for a better sustainable management.
This study is carried out in the firstly in the herbaria of the University Cheikh Anta Diop of Dakar/Senegal: the herbarium
of the Herbarium of the Institut Fondamental d'Afrique Noire (IFAN UCAD) and the herbarium DAKAR of the
Department of Plant Biology of the Faculty of Sciences and Techniques. Secondly, a consultation of different checklists
and databases was conducted to collect more information. This research work on the flora of liverworts is rich with
nearly 39 species divided into 24 genera and 13 families. The order Jungermanniales is dominant in terms of families
and the Porellales Jubulineae in terms of genera and species. The family Lejeuneaceae is in the majority in this flora
before the families Ricciaceae, Plagiochilaceae and Metzgeriaceae.
These results are important for the conservation of the environment and the development of the plant biodiversity of
Senegal.
Keywords: Liverworts; IFAN herbarium; DAKAR herbarium; Biodiversity; Senegal
1. Introduction
The Marchantiophyta comprise about 5000-6000 species in 391 genera [26, 7, 14, 19, 20, 15, 8, 9, 35] sometimes even
reaching 7500 species [37, 38] of which 80% are leafy liverworts. According to the results of recent monographic work,
these species show a cosmopolitan distribution [33, 34, 32, 7, 13]. The Marchantiophyta are the earliest diverging
lineage in terrestrial plants, probably dating from the Silurian period [25, 31, 36, 39, 17, 18]. In Senegal, studies on the
specific diversity of liverworts are still at an embryonic stage, despite the presence of samples in the Dakar and IFAN
herbaria.
The present work aims to contribute to a better knowledge of the diversity of liverworts in the herbaria of Senegal in
order to provide a data base for a sustainable management.
Corresponding author: Djibril Diop
Botany - Biodiversity Laboratory, Department of Plant Biology, Faculty of Science and Technology, Cheikh Anta Diop University, BP
5005, Dakar-Fann, Senegal.
Copyright © 2021 Author(s) retain the copyright of this article. This article is published under the terms of the Creative Commons Attribution Liscense 4.0.
GSC Biological and Pharmaceutical Sciences, 2021, 17(03), 078–084
2. Material and methods
2.1. Study sites
2.1.1. The Herbarium of the Institut Fondamental d'Afrique Noire (IFAN)
The Herbarium of the Institut Fondamental d'Afrique Noire (IFAN UCAD) created in 1941 by Professor Paul JAEGER is
the oldest herbarium in French-speaking Africa. Indeed, it centralized most of the botanical activities in the former
French metropolis. Its main objective was to create a bank of all the plant species of French-speaking Africa, then of all
continental and insular Africa with nearly 60,000 specimens.
2.1.2. The Herbarium of the Department of Plant Biology: DAKAR Herbarium
Created in 1960 by Professor J. MIEGE and rehabilitated in 1994, the DAKAR herbarium is the second herbarium of
Senegal. The main objective is to constitute a collection of good quality and representative of the flora of Senegal and a
remarkable knowledge base in the fields of research, training, knowledge, conservation of plant resources and African
and international collaboration. The herbarium houses about 13000 samples.
2.2. Material
The material consisted of samples from the DAKAR and IFAN herbaria and samples reported in the Checklist and
distribution of the liverworts and hornworts of Sub-Saharan Africa, including the East African Islands [41].
Consultations of herbarium collections and databases were carried out to establish the floristic list of liverworts.
2.3. Methods
Scientific names were verified via Tropicos, Plant List and efloras. The classification of [35]; was used for liverworts.
African identification keys: for liverworts [40], from [22, 23], the flora of Kenyan mosses and liverworts [4]; volume 14
of the Rwandan Liverworts and Hornworts [11] and Catalogue of the Hepaticae and Anthocerotae of sub-Saharan Africa
[42] were used to facilitate identification as well as Augier's [2, 3] flora of Bryophytes and many other articles.
3. Results
3.1. Specific composition
Table 1 represents the global list of species and families encountered during this study. Thus 39 species of liverworts
were recorded in these herbaria and check-list belonging to 7 orders, 24 genera and 13 families.
3.2. Taxonomic spectrum
Table 2 gives indications on the structure of the Marchantiophyta flora. These results show a great specific diversity of
these herbarium collections.
The order Jungermanniales are dominant in terms of families with 38.5% and the Porellales Jubulineae in terms of genera
with 54.2% and species with 43.6%. Except for the order Metzgeriales with 2 families; all other orders are represented
by a single family.
The analysis of Table 3 of the distribution of genera and species in the families shows an important diversity. This flora
is dominated by species of the family Lejeuneaceae occupying 50% of the genera and 41% of the species of the total
number. All other remaining families are represented by only one genus. In terms of species, after the Lejeuneaceae
come the species of the families Ricciaceae, Plagiochilaceae and Metzgeriaceae with 15%, 13% and 7.7% respectively. It
appears from this analysis that the remaining families are represented by only one species.
79
GSC Biological and Pharmaceutical Sciences, 2021, 17(03), 078–084
Table 1 List of species of Marchantiophyta surveyed
Orders
Metzgeriales Roseuv.
Families
Species
Aneuraceae H. Klinggr.
*Riccardia limbata (Stephani) E.W. Jones
*Metzgeria australis Stephani
Metzgeriaceae H. Klinggr.
*Metzgeria camerunensis Stephani
*Metzgeria hamata Lindb.
Marchantiales Limpr.
Dumortieraceae D.G. Long
*Dumortiera hirsuta (Sw.) Nees
*Riccia discolor Lehm. & Lindenb.
R. intermedia E.W. Jones
*Riccia frostii Austin
Ricciales L.
Ricciaceae Rchb.
*Riccia fluitans L.
*Riccia cavernosa Hoffm.
R. rautanenii (Steph.) Steph.
***Riccia sp1
***Riccia sp2
Herbertaceae Müll. Frib. Ex *Herbertus juniperoideus (Sw.) Grolle. Herbertia juniperina
Fulford et Hatcher
(Sw.) Evs.
Lepidoziaceae Limpr.
*Bazzania herminieri (Gottsche ex Steph.) Pagán
Bazzania herminieri (Gottsche)
Lophocoleaceae Müll. Frib.
*Lophocolea difformis Nees
ex Vanden Berghen
Jungermanniales
H.
Klinggr. - Lophocoleineae
Schljakov.
*Plagiochila barteri Mitt.
*Plagiochila pectinata Lindenb.
P. pectinata Wild ex Lindenb
Plagiochilaceae Müll. Frib.
*Plagiochila lastii Mitt.
Plagiochila terebrans Nees et Mont. Lindenb.
*Plagiochila sp1
***Plagiochila sp 2
Trichocoleaceae Nakai
*Trichocolea tomentosa (Sw.) Nees
Jubulaceae H. Klingger.
*Frullania serrata Gottsche.
*Odontolejeunea tortuosa (Lelm. et Lindenb.) Stephani
*Cololejeunea obliqua (Nees et Mont.) Schiffner
*Cololejeunea leloutrei (E.W. Jones) R.M. Schust.
*Diplasiolejeunea cavifolia Stephani
*Hygrolejeunea acuta (Mitt.) Vander Bergham
*Leptocolea cuneifolia (Stephani) A. Evans
Porellales
Schljakov.
Jubulineae Müll. Frib.
Lejeuneaceae Casares-Gil
*Leptocolea sp
**Acrolejeunea emergens (Mitt.) Steph. var. confertissima
(Steph.) Gradst.
**Acrolejeunea emergens (Mitt.) Steph. var. emergens
*Ceratolejeunea cornuta (Lindenb.) Steph.
C. jungneri Steph.
*Drepanolejeunea physifolia (Gottsche) Steph.
D. clavicornis Steph.)
*Drepanolejeunea cultrella (Mitt.) Stephani
80
GSC Biological and Pharmaceutical Sciences, 2021, 17(03), 078–084
*Lejeunea flava (Sw.) Nees
*Leptolejeunea truncatiflora Steph.
*Prionolejeunea grata (Gottsche) Schiffn
P. serrula (Mitt.) Stephani;
P. kindbergii (Steph.) Schiffn
**Thysananthus auriculatus (Wilson et Hook.) Sukkharak et
Gradst. Mastigolejeunea auriculata (Wilson et Hook.) Steph.
Mastigolejeunea auriculata (Wilson) Schiffn. (Steph.)
Gradst.)
Porellales
Schljakov.
Radulaceae Müll. Frib.
Radulineae R.M. Schust.
*Radula flaccida Lindenb. & Gottsche.
Pleuroziales Schljakov.
*Pleurozia gigantea (F. Weber) Lindb.
Pleuroziaceae Müll. Frib.
*= IFAN Herbarium, **= Check-List of Wigginton, ***= DAKAR Herbarium
Table 2 Structure of the Marchantiophyta Flora
Ordres
Families
Genera
Species
Nomber
%
Nomber
%
Nomber
%
Porellales Jubulineae
2
15,4
13
54,2
17
43,6
Jungermanniales
5
38,5
5
20,8
9
23,1
Metzgeriales
2
15,4
2
8,3
4
10,3
Marchantiales
1
7,7
1
4,2
1
2,6
Ricciales
1
7,7
1
4,2
6
15,4
Porellales Radulinea
1
7,7
1
4,2
1
2,6
Pleuroziales
1
7,7
1
4,2
1
2,6
Total
13
100
24
100
39
100
Table 3 Distribution of species in the different taxonomic groups
Families
Genera
Species
Nomber
%
Nomber
%
Lejeuneaceae
12
50
16
41
Ricciaceae
1
4,2
6
15
Plagiochilaceae
1
4,2
5
13
Metzgeriaceae
1
4,2
3
7,7
Aneuraceae
1
4,2
1
2,6
Dumortieraceae
1
4,2
1
2,6
Herbertaceae
1
4,2
1
2,6
Lepidoziaceae
1
4,2
1
2,6
Lophocoleaceae
1
4,2
1
2,6
Trichocoleaceae
1
4,2
1
2,6
Jubulaceae
1
4,2
1
2,6
Radulaceae
1
4,2
1
2,6
Pleuroziaceae
1
4,2
1
2,6
24
100
39
100
Total
81
GSC Biological and Pharmaceutical Sciences, 2021, 17(03), 078–084
4. Discussion
This study shows that the flora of liverworts in the herbaria and checklists is rich with nearly 39 species distributed in
24 genera and 13 families. This richness is explained by the fact that Senegal was a strategic point in the management
of herbarium collections in Black Africa and the world. The IFAN’s herbarium has hosted during missions collections of
bryophytes and phanerogams of several authors.
Moreover, the collections of Marchantiophytes of this herbarium represent almost a great part of this flora. Compared
to the liverwort flora of Madagascar [28] or the Reunion Islands [1], the latter is less diversified because there are very
few field studies, contrary to other countries where many investigations have been conducted for the knowledge of the
bryological flora.
At the level of families, the dominance of the family Lejeuneaceae is a characteristic of the flora of liverworts in the
tropical zone. The Lejeuneaceae are the largest family of the liverworts (Hepaticae), with almost a thousand species in
91 currently accepted genera. They abound in humid tropical forests where they inhabit a large variety of niches,
including tree trunks, branches, saplings, twigs and the surfaces of living leaves. In an average tropical lowland rain
forest, more than three-quarters of the hepatic species are Lejeuneaceae [12, 16].
The Ricciaceae form a cosmopolitan family with two genera (Riccia, Ricciocarpos). The genus Riccia is represented by
about 200 species worldwide [43], including 88 for sub-Sahelian Africa, with the Indian Ocean [40]. Only the Riccia of
Mediterranean North Africa [24] and South Africa have been explored and described in some detail [29]. Riccia are very
diverse in South Africa (about 80 species), and they show a high rate of endemism, rare for bryophytes.
Compared to other African floras, this one is similar with a strong representation of mosses on liverworts: Guinea
Conakry [27, 41] and African floras: Reunion Island [1], Mauritius [21, 10], South Africa [30], the Kenyan mosses and
liverworts flora [5, 6].
5. Conclusion
The aim of the present work is to contribute to a better knowledge of liverworts in the Senegalese herbaria in order to
provide basic knowledge for a better sustainable management. This study shows that the flora of liverworts in the
herbaria and check-list is rich with nearly 39 species distributed in 24 genera and 13 families. The order
Jungermanniales is dominant in terms of families and the Porellales Jubulineae in terms of genera and species. This flora
is dominated by species of the family Lejeuneaceae occupying 50% of the genera and 41% of the species of the total
number. In terms of species, after the Lejeuneaceae come the species of the family Ricciaceae, Plagiochilaceae and
Metzgeriaceae. It also appears from this analysis that the remaining families are represented by only one species.
Compliance with ethical standards
Acknowledgments
Our thanks to the IFAN and DAKAR herbaria and to the researchers and technicians for their help and availability.
Disclosure of conflict of interest
The authors declare no conflict of interest.
References
[1]
Ah-Peng C, Diversity, distribution and biogeography of bryophytes from Lava flows of the Piton de la Fournaise
volcano (La Réunion, France). PhD University of La Réunion, Saint-Denis. 2007.
[2]
Augier, J. Flora of bryophytes. Paris. 1966.
[3]
Augier J. Keys of determination for the Hepatics and Anthocerota in Cameroon forest. I. [1]-123. Text, Tables,
Glossary. II. [124]-155. Plates, Legends. Paris, the author. [Contribution to the Flora of Cameroon (Cryptogamy)
–an essay for students and researchers in the RUC an d neighboring W and Central African regions]. 1985.
82
GSC Biological and Pharmaceutical Sciences, 2021, 17(03), 078–084
[4]
Chuah-Petiot MS. Contribution to the bryoflora of Kenya. Tropical Bryology. 2001; 20: 31-37.
[5]
Chuah-Petiot MS. Mosses, Liverworts and Hornworts of Kenya p. 273. An illustrated guide with descriptions and
figures of over 300 species and keys for identification. Edited by the author. 2003.
[6]
Chuah-Petiot MS, Pócs T. East African bryophytes XIX. A contribution to the bryoflora of Kenya. Acta Botanica
Hungarica. 2003; 45: 53-64.
[7]
Crandall-Stotler B. Stotler RE. Morphology and classification of the Marchantiophyta. In A. J. Shaw and B. Goffinet
[eds.], Bryophyte biology. Cambridge University Press, Cambridge, UK. 2000; 21–70.
[8]
Crandall-Stotler BJ, Forrest LL, Stotler RE. Evolutionary trends in the simple thalloid liverworts
(Marchantiophyta, Jungermanniopsida subclass Metzgeriidae). Taxon. 2005; 54: 299–316.
[9]
Crandall-Stotler BJ, Stotler RE, Long DG. Morphology and classification of the Marchantiophyta. In: Goffinet, B. &
Shaw, A. J. (eds) Bryophyte Biology, 2nd edition, pp. 1–54. Cambridge: Cambridge University Press. 2008.
[10]
Een G. Bryophytes collected in Mauritius by Gunnar Erdtman. Tropical Bryology. 2000; 18: 87-90.
[11]
Fischer E. Liverworts and Hornworts of Rwanda. Abc Taxa. 2013; 14: 552.
[12]
Gradstein SR. Lejeuneaceae: Ptychantheae, Brachiolejeuneae. Flora Neotropica Monograph. 1994; 62(1): 216.
[13]
Gradstein SR, Churchill SP. Salazar A.N. Guide to the Bryophytes of Tropical America. Memoirs of the New York
Botanical Garden. 2001; 86: 577.
[14]
Gradstein SR, Costa DP. The liverworts and hornworts of Brazil. Memoirs of the New York botanical garden. 2003;
87: 1–317.
[15]
Gradstein SR, Ilkiu-Borges AL. Guide to the plants of Central French Guiana. Part IV. Liverworts and hornworts.
Memoirs of the New York botanical garden. 2009; 76(4): 1–140.
[16]
Gradstein SR, Pócs T, Zhu RL. Lejeuneaceae. In: Söderström, L., Hagborg, A. & von Konrat, M. (Eds.) World
checklist of hornworts and liverworts. PhytoKeys. 2016; 59: 406–408.
[17]
Groth-Malonek M, Knoop V. Bryophytes and other basal plants: the mitochondrial perspective. Taxon. 2005.
[18]
Groth-Malonek M, Pruchner D, Grewe F, Knoop V. Ancestors of Trans-Splicing Mitochondrial Introns Support
Serial Sister Group Relationships of Hornworts and Mosses with Vascular Plants Article in Molecular Biology and
Evolution. 2005.
[19]
Heinrichs J, Hentschel J, Wilson R, Feldberg K, Schneider H. Evolution of leafy liverworts (Jungermannniidae,
Marchantiophyta): Estimating divergence times from chloroplast DNA sequences using penalized likelihood with
integrated fossil evidence. Taxon. 2007; 56: 31–44.
[20]
Heinrichs J, Hentschel J, Feldberg K, Bombosch A, Schneider H. Phylogenetic biogeography and taxonomy of
disjunctly distributed bryophytes. Journal of Systematics and Evolution. 2009; 47(5): 497-508.
[21]
Johnston HH. Additions to the flora of Mauritius, as recorded in Baker's "Flora of Mauritius and the Seychelles".
Transactions and Proceedings of the Botanical Society of Edinburgh. 1895; 59: 391-407.
[22]
Jones EW. African Hepatics XII. Some little-known Lejeuneaceae. Transactions of the British Bryological Society.
1975; 3: 191-207.
[23]
Jones EW. African Hepatics XIII. The Ricciaceae of Tropical Africa. Transactions of the British Bryological Society.
1957; 3: 208-227.
[24]
Jovet-Ast S. Towards a phylogenetic classification of species in the genus Riccia. The Bryologist. 90(4): 321–330.
1987.
[25]
Kenrick P, Crane PR. The origin and early evolution of plants on land. Nature. 1997; 389 (6646): 33-39.
[26]
Lewis L, Mishler BD, Vilgalys R. Phylogenetic relationships of the liverworts (Hepaticae), a basal embryophyte
lineage, inferred from nucleotide sequence data of the chloroplast gene rbcL. Molecular Phylogenetics and
Evolution 7: 377–393. 1997.
[27]
O’Shea BJ. Checklist of the mosses of sub-Saharan Africa (version 5, December 2006). Tropical Bryology Research
Reports. 2006; 6: 1.255.
[28]
Marline L, Andriamiarisoa RL, Bardat J, Chuah-Petiot M, Hedderson TAJ, Reeb C, Strasberg D, Wilding N, Ah-Peng
C. Checklist of the bryophytes of Madagascar. Cryptogamie, Bryologie. 2012; 33: 199-255.
83
GSC Biological and Pharmaceutical Sciences, 2021, 17(03), 078–084
[29]
Perold SM. Studies in the genus Riccia (Marchantiales) from southern Africa. 7. Riccia congoana and its synonyms. Bothalia. 1986; 16: 193-201.
[30]
Perold SM. Hepatophyta Part 1. Marchantiales. Fasc.1. Marchantiidae. Flora of Southern Africa (ed. by O.A.
Leistner). National Botanical Institute, Pretoria. 1999.
[31]
Qiu YL, Cho Y, Cox JC, Palmer JD. The gain of three mitochondrial introns identifies liverworts as the earliest land
plants. Nature. 1998; 394: 671–674.
[32]
Schuster RM. Evolution, phylogeny and classification of the Hepaticae. 1070 Pages 892–In R.M. Schuster (ed.),
New Manual of Bryology. Hattori Botanical Laboratory, Nichinan, Japan. 1984.
[33]
Schljakov RN. On the higher taxa of liverworts – class Hepaticae s. str. [Bot. Zhurn. (Moscow & Leningrad): Names
and titles transliterated and translated from the Russian; In Russian, English summary]. 1972; 57: 496–508.
[34]
Schljakov RN. Hepatics: Morphology, Phylogeny, and Classification. [Leningrad: Russian Acad. Sci.] [Names and
titles transliterated and translated from the Russian; In Russian.] 1975.
[35]
Söderström L, Hagborg A, von Konrat M, Bartholomew-Began S, Bell D, Briscoe L, Brown E, Cargill DC, Costa DP,
Crandall-Stotler BJ, Cooper ED, Dauphin G, Engel JJ, Feldberg K, Glenny D, Gradstein SR, He XL, Heinrichs J,
Hentschel J, Ilkiu-Borges AL, Katagiri T, Konstantinova NA, Larrain J, Long DG, Nebel M, Pócs T, Felisa-Puche F,
Reiner-Drehwald E, Renner MAM, Sass-Gyarmati A, Schafer-Verwimp A, Segarra- Moragues JG, Stotler RE,
Sukkharak P, Thiers BM, Uribe J, Váňa J, Villarreal JC, Wigginton M, Zhang L, Zhu RL. World checklist of hornworts
and liverworts. PhytoKeys. 2016; 59: 1.828.
[36]
Steinhauser S, Beckert S, Capesius I, Malek O, Knoop V. Plant mitochondrial RNA editing. Journal of Molecular
Evolution. 1999; 48: 303–312.
[37]
von Konrat MJ, Renner M, Söderström S, Hagborg A, Mutke J. Early Land Plants Today: Liverwort species diversity
and the relationship with higher taxonomy and higher Plants. Fieldiana, Botany. 2008; 2(193): 87–102.
[38]
von Konrat MJ, Söderström S, Hagborg A. The Early Land Plants Today project (ELPT): A communitydriven effort
and a new partnership with Phytotaxa. Phytotaxa. 2010; 9: 11–21.
[39]
Wellman CH, Osterloff P, Mohluddin U. Fragments of the earliest land plants. Nature. 2003; 425: 282–285.
[40]
Wigginton MJ. Checklist and distribution of the liverworts and hornworts of Sub-Saharan Africa, including The
East African Island Tropical. Bryology Research Report. 2004; 5: 1–102.
[41]
Wigginton MJ. Checklist and distribution of the liverworts and hornworts of Sub-Saharan Africa, including the
East African Islands. Edition 3. Tropical Bryology Research Reports. 2009; 7: 1. 114.
[42]
Wigginton MJ, Grolle R. Catalogue of the Hepaticae and Anthocerotae of sub-Saharan Africa. Bryophyterum
Bibliotheca. 1996; 50: 1-267.
[43]
Wheeler JA. Molecular phylogenetic reconstructions of the Marchantioid liverworts radiation. The Bryologist.
2000; 103: 314–333.
84