Corresponding author: Komla Elikplim Abotsi (
Academic editor: Daniele Cicuzza
The
This study provides first local scientific information about the fern flora of Togo. It confirmed the presence of
Ferns have been studied, in the past century, mainly using morphological traits whereas more light have been brought to the group with the recent development of cytology and molecular phylogeny (
As a large family of leptosporangiate ferns (about 10% of the Monilophytes),
The
The family is well present in Togo but few studies hitherto focused on it. The current data of the country's biodiversity (
Recent revisions of the family are not taken into account neither at the herbarium of the University of Lomé nor in the flora of Togo. Many names have then been placed in synonymy in recent databases (
The current study aims at updating the
The study area is Togo. It is located in the Gulf of Guinea, and surface covers approximately 56 785 km2. Its forest surface is counts among the least in the West African sub-region.
Three main types of climate are found in Togo. In the North, there is the tropical climate with variations increasingly wet, going to the South. The southern coastal area has a sub-equatorial climate. A transitional climate between these two types is found in the mountains of the south. Togo has varied soils. There are five main soil classes (
There have been new harvest in the ecologic zone 4 of Togo in order to enrich the plant materials available for this study.
The study is based on new harvests from the ecological zone 4 (215 specimens) and specimens from herbaria of the University of Lomé (TOGO) (87 specimens) and the National Museum of Natural History in Paris (P) (122 specimens).
New harvests were conducted along the banks of the natural ecosystem's streams in the study area along a topographic gradient. As many others authors,
Data from this new inventory is coupled with ecological and geographical data from previously performed harvests between 1973 and 1994 across the country and available in the RIHA data base (African Herbaria Computer Network) of the herbarium of Lomé.
Each harvest consists of one or more photos of the whole plant in its habitat, a herbarium specimen represented by the whole plant (or a fragment then comprising parts of all organs) and a piece of lamina in silica gel (
The morphological observations of all available specimens (new specimens and old herbarium collections) enabled the establishment of a list of morphological descriptors which have been added to the ecological characteristics of each species (Table
Xper² is a free software for management, editing, and analysis of taxonomic knowledge bases (
All species identified as a result of the study and the identification of all available specimens were then associated with the states of the descriptors that characterize them. This same approach was also used by
The different states of retained descriptors were used to create a dichotomous identification key of the Togolese
Geographic data were collected from herbarium specimen labels and new harvests sheets in order to edit a distribution map of the
Native
Least Concern
Togo (Ecological Zone 5), USA, U.S. Virgin Isl., Costa Rica, Galapagos, Puerto Rico, Hispaniola, Jamaica, Cuba, Nicaragua, Barbados, Mexico, Belize, Guatemala, Honduras, Panama, Colombia, Venezuela, Ecuador, Peru, Brazil, Trinidad, Guadeloupe, Martinique, St. Lucia, Grenada, Barbados, Guyana, Surinam, French Guiana, Cayman Isl., Bahamas, Turks & Caicos Isl., Isla del Coco, Australia, South Africa, Zambia, Mozambique, SE-Zimbabwe, Kenya, Tanzania, Zanzibar, Pemba Isl., Principe Isl., Sao Tomé, Bioko Isl., Senegal, Guinea-Bissau, Guinea, Gambia, Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Equatorial Guinea, Gabon, Angola, Madagascar, Mauritius, La Réunion, Micronesia, Palau Isl., Fiji, Western Samoa, American Samoa, Marianas, Society Isl., Tonga, New Caledonia, Niue, Austral Isl., China, Taiwan, Ryukyu Isl., peninsular Malaysia, India, Andaman Isl., Nicobar Isl., Philippines, Sri Lanka, Vietnam, Thailand, Cambodia, Sulawesi
Native
Togo (Ecological Zone 2), South Africa, Namibia, Swaziland, Botswana, Zimbabwe, Mozambique, Zambia, D.R.Congo, Zimbabwe, Uganda, Kenya, Tanzania, Zanzibar, Djibouti, Mali, Nigeria, Cameroon, Somalia, Madagascar, Comores, Seychelles, La Réunion, Mauritius, Angola, Sudan, SE-Egypt, Ethiopia, N-Yemen, Saudi Arabia, India, Sri Lanka, Iran, Afghanistan, Pakistan, Nepal, Burma, Cape Verde Isl.
Small fern with an erect rhizome and fronds in clumps, with lanceolate scales toothed at their base,
Native
Togo (Ecological Zone 4), South Africa, Swaziland, Namibia, Zimbabwe, Zambia, Malawi, Mozambique, Ghana, Nigeria, Uganda, Kenya, Tanzania, Somalia, Madagascar, Ivory Coast, Angola, Ethiopia, N-Yemen, S-Yemen, Oman, Saudi Arabia, Pakistan, Galapagos, Jammu & Kashmir, India, Cape Verde Isl.
Native
Togo (Ecological Zones 3 and 4), China, Taiwan, Australia, Jammu & Kashmir, India, Andaman Isl., Myanmar, Philippines, Thailand, Laos, Cambodia, Vietnam, peninsular Malaysia, Sulawesi, Sri Lanka, Nepal, Moluccas, Oman, S-Yemen, Zimbabwe, Mozambique, South Africa, Zambia, Malawi, Chad, Tanzania, Sao Tomé, Bioko Isl., Senegal, Guinea-Bissau, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Benin, Niger, Nigeria, Cameroon, Central African Republic, Sao Tome, Congo, D.R.Congo, Angola, Ethiopia, Sudan, Madagascar, Comores, Burkina Faso, Mali, Sudan, Southern Marianas, Fiji, Micronesia, Palau Isl., Western Samoa, Cuba, Cape Verde Isl.
Native
Togo (Ecological Zones 2 and 4), Tanzania, Chad, Mali, Senegal, Sudan, Central African Republic, Guinea, Nigeria, D.R.Congo, Angola, Cameroon
Native
Togo (Ecological Zones 4 and 5), Uganda, Zanzibar, Principe Isl., Bioko Isl., Senegal, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Benin, Nigeria, Cameroon, Central African Republic, Equatorial Guinea, Gabon, Congo, D.R.Congo, Angola
Native
Least Concern
Togo (Ecological Zones 2, 3 and 5), USA, Costa Rica, Nicaragua, Mexico, Guatemala, El Salvador, Panama, Colombia, Venezuela, Puerto Rico, Jamaica, Hispaniola, Ecuador, Brazil, Guyana, Surinam, French Guiana, Australia, Taiwan, China, Japan, Ryukyu Isl., India, Andaman Isl., Nicobar Isl., Sri Lanka, Myanmar, Nepal, Philippines, Vietnam, Thailand, Laos, Cambodia, Sulawesi, Hawaii (I) (Kauai (I), Oahu (I)), Fiji (I), Micronesia, Palau Isl., Southern Marianas, Uganda, Tanzania, Kenya, Zanzibar, Somalia, Sudan, Congo, Angola, Zambia, Zimbabwe, Mozambique, Ethiopia
Native
Togo (Ecological Zones 3 and 4), Zambia, Zimbabwe, Malawi, Tanzania, D.R.Congo, Madagascar, Ghana, Nigeria, Central African Republic
Native
Togo (Ecological Zone 4), South Africa, Namibia, Swaziland, Botswana, Zambia, Zimbabwe, Malawi, Mozambique, Angola, Cameroon, Central African Republic, Liberia, Tanzania, Uganda, Madagascar, Comores, Guinea, Ivory Coast, Ghana, Nigeria, Socotra, Ethiopia, S-India, Sri Lanka, Philippines, Vietnam, Sulawesi, Fiji, Society Isl., Tonga, New Caledonia, Western Samoa, American Samoa, Marquesas Isl., Austral Isl., Australia, Taiwan, China
Native
Togo (Ecological Zone 4), Congo, Guinea, Sao Tomé, Principe Isl., Bioko Isl., Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Central African Republic, Equatorial Guinea, Gabon, Congo, Angola, Uganda
Native
Togo (Ecological Zones 3 and 4), Burundi, Ghana, Malawi, Tanzania, Uganda, Zambia, Kenya, Mozambique, Seychelles, Principe Isl., Sao Tomé, Chad, Senegal, Guinea, Sierra Leone, Liberia, Ivory Coast, Benin, Bioko Isl., Nigeria, Cameroon, Central African Republic, Annobon Isl., Congo, D.R.Congo, Angola, Mali, Sudan
Native
Togo (Ecological Zones 2 and 4), Bahamas, Mexico, Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Paraguay, Uruguay, St. Martin, St. Barthelemy, Antigua, Saba, St. Eustatius, St. Kitts, Montserrat, Guadeloupe, Marie Galante, Dominica, Martinique, St. Lucia, St. Vincent, Grenada, Barbados, Guyana, Surinam, French Guiana, Puerto Rico, Cuba, Jamaica, Hispaniola, Vieques Isl., Virgin Isl., Curacao, Isla del Coco, Australia (I), Taiwan (I), Ryukyu Isl. (I), China (I), Mozambique (I), Gabon (I), Uganda (I), Tanzania (I), Bioko Isl. (I), Sao Tomé and Principe (I), Senegal (I), Guinea (I), Sierra Leone (I), Liberia (I), Ivory Coast (I), Ghana (I), Nigeria (I), Cameroon (I), Gabon (I), Congo (I), Angola (I), D.R.Congo, Zambia (I), Madagascar (I), Mauritius (I), La Réunion (I), Comores (I), Seychelles (I), Burkina Faso (I), Marquesas Isl. (I), Society Isl. (I), Southern Marianas (I), Fiji (I), Palau Isl. (I), Micronesia (I), Hawaii (I), Malaysia (I), Borneo (I), Thailand (I), India (I), Sri Lanka (I), Andaman Isl. (I), Nicobar Isl. (I), Cambodia (I), Vietnam (I), Laos (I), Philippines (I), Sulawesi (I), Moluccas (I), USA (I)
Native
Togo (Ecological Zone 4), Gabon, Comores, Uganda, Kenya, Tanzania, Pemba Isl., Principe Isl., Sao Tomé, Bioko Isl., Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Benin, Nigeria, Cameroon, Central African Republic, Equatorial Guinea, Congo, D.R.Congo, Angola, Burundi, Sudan, Seychelles
The rhizome of
Native
Togo (Ecological Zones 2, 4 and 5), Guinea, Sierra Leone, Tanzania, Principe Isl., Bioko Isl., Ivory Coast, Liberia, Ghana, Nigeria, Cameroon, Central African Republic, Gabon, Congo, D.R.Congo, Angola, Burundi
The rhizome of
Native
Togo (Ecological Zone 4), Uganda, Tanzania, Principe Isl., Sao Tomé, Bioko Isl., Guinea, Mali, Sierra Leone, Liberia, Ivory Coast, Ghana, Benin, Nigeria, Cameroon, Central African Republic, Gabon, Congo, D.R.Congo, Angola, Sudan
Native
Togo (Ecological Zones 2, 3, 4 and 5), Principe Isl., Bioko Isl., Senegal, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Central African Republic, Gabon, Congo, Angola, Sudan, Madagascar, Mali
Native
Togo (Ecological Zone 4), Australia, Taiwan, China, Singapore, Borneo, S-India, Andaman Isl., Nicobar Isl., Sri Lanka, peninsular Malaysia, Philippines, Thailand, Sulawesi, Sumatra, Laos, Nepal, Vietnam, Java, Moluccas, Uganda, Kenya, Tanzania, Zanzibar, Principe Isl., Sao Tomé, Chagos Arch., Galega Isl., Bioko Isl., Guinea, Sierra Leone, Ivory Coast, Ghana, Benin, Liberia, Nigeria, Cameroon, Central African Republic, Equatorial Guinea, Gabon, Congo, Madagascar, La Réunion, Comores, Seychelles, Mauritius, Angola, Marianas, Fiji, Palau Isl., Micronesia, Marshall Isl., Nauru, Gilbert Isl., Tonga, New Caledonia, American Samoa, Western Samoa, Tuvalu, Society Isl., Austral Isl., Niue, Marquesas Isl., Costa Rica (I), Costa Rica (I), Panama (I), Colombia (I), Venezuela (I), Brazil (I), Ecuador (I), Peru (I), St. Lucia (I), Puerto Rico (I), Jamaica (I), Surinam (I), French Guiana (I), Mexico (I), USA (I)
1 | Aquatic plant, partially or totally submerged |
|
– | Terrestrial plant, epiphytic, or lithophyte |
|
2 | Mangrove plant, lamina coriaceous, oblong-lanceolate, pinnate, lower pinnae short-stalked and terminal pinnae sessile, acrostichoids sori (sporangia covering completely the lower surface of the lamina) |
|
– | Fresh water plant, completely submerged, lamina membranous, lanceolate, fertile frond, 2-4 pinnate, sporangia protected by the reflected margin of lamina |
|
3 | Frond entire or flabellate |
|
– | Frond pinnate, pinnatifid, multi-pinnate or multi-pinnatifid |
|
4 | Plant epilithic, lamina fan-shaped, flabellate, coriaceous, stipe sparsely scaled |
|
– | Epiphytic plant, lamina linear-lanceolate, stipe blackish at the base, marginal sori immersed |
|
5 | Frond pinnate |
|
– | Frond multi-pinnate or multi-pinnatifid |
|
6 | Pinnae long-stalked |
|
– | Pinnae sessile or sub-sessile |
|
7 | Pinnae deeply or slightly incised, generally hairy, becoming smaller at the top of the leaf, blade oblong to deltoid, rachis and stipe dark brown, densely hairy |
|
– | Pinnae entire or lobed, oblong, oval or lanceolate, glabrous, stipe and rachis black and shiny |
|
8 | Sori linear, protected by the reflected margin of lamina, terminal pinnae similar to lateral ones, lamina dark green, with coriaceous texture |
|
– | Sori oblong-kidney shaped, pinnae lobed and serrated feathers on the upper margin, light green leaf blade, papery textured lamina |
|
9 | Sori moon crescent shaped, rhomboid-dimidiate pinnae, lamina upper margin incised |
|
– | Sori linear, pinnule rhomboidal, non-dimidiate |
|
10 | Stipe black |
|
– | Stipe not black |
|
11 | Limb oblong-lanceolate, not winged, covered by a white powdery layer on the underside |
|
– | Limb deltoid, broadly winged, 3-branched at the base, sori marginal |
|
12 | Sori linear, interrupted |
|
– | Sori linear, continuous over the entire margin of the lamina, except at the base of sinus |
|
13 | Limb oval-deltoid light green, 3-pinnate, equal pinnae, lamina herbaceous, stipe tri-branched at the base of the limb |
|
– | Limb lanceolate to deltoid, oblong, elliptic or oval, basal pinnae developed basiscopically, stipe not branched |
|
14 | Sori linear or not reaching the bottom of the base or the apex of the pinnae, stipe reddish at the base |
|
– | Sori U-shaped, interrupted at the top of pinnae |
|
15 | Frond deltoid, more or less lanceolate, brown stipe, simple or pinnatifid pinnae, both types can coexist on the same frond, oblong-lanceolate |
|
– | Frond not deltoid, stipe straw colored, pinnatifid pinnae only |
|
16 | Stipe channeled, muricate, purplish or brownish at the base, pinnae and pinnule oblong-lanceolate, lamina proliferous |
|
– | Stipe reddish at the base, pinnae oblong, pinnule oblong-falcate, limb non-proliferous |
|
From all the studied specimens, 17 species have been identified, belonging to the
The recognition and distinction of taxa within the
Most of recorded species (14 species of 17 in total) are represented in the ecological zone 4. Some among them are unique to this area (
Three families in the RIHA database of the herbarium of Lomé correspond to the current delimitation of the
Togo ecological zone 4 concentrates 94.12 % of the species diversity of Togolese
The expansion of croplands is also a real threat to the
Similarly, the current prerogatives on the carbon stock promote species that store more carbon without necessarily referring to the local flora. Therefore, timber plantations like
This study clarifies the nomenclature and systematic boundaries of the
This study is a basis for future valorization of the family. Considerations should be given to
Togo ecological zones.
Ecological Zone 1: Refers to the Northern Plains. It enjoys a Sudanese climate characterized by an average of 6 to 7 months of drought. Rainfall varies from 800 to 1000 mm and water per year. The temperatures are between 17 and 39° C. The predominant vegetation is the Sudan savanna with few islands of dry forests and gallery forests. The soil is tropical ferruginous, sandy.Ecological Zone 2: It is covered with a mosaic of dry forests of mountain and forest galleries. The climate is Sudano-Guinean. The rainy season extends from April to October and the dry season is characterized by the presence of the Harmattan, a dry and cold wind. Rainfall is irregular and reaches 1200-1300 mm of water per year. The soils are thin and contain a high proportion of coarse elements. Ferruginous tropical soils are also present.Ecological Zone 3: It corresponds to the Guinean savannas of central area plains enjoying a tropical climate with a rainy season extends from May to October. Rainfall varies between 1200 and 1500 mm per year. The temperature is between 25 and 40° C. The soils are mainly ferruginous. Semi-deciduous forests are found in the southern part and dry forests in the northern part.Ecological Zone 4: This corresponds to the southern part of the Mount "Togo" and has a sub-equatorial climate of transition, characterized by a long rainy season from March to October with a small interruption in August or September. Rainfall varies between 1300 and 1600 mm per year. The vegetation is constitute of rainforests, on deep red lateritic soils.Ecological Zone 5: It is the South of the coastal plains area, with a sub-equatorial climate, characterized by a lack of rainfall in it's southern part. Rainfall varies from 800 mm at the coast up to 1200 mm per year at the northern limit of the zone. The vegetation is composed by a mosaic of savannah, farmland and dry forests (
Species diversity by gender within
Distribution of
General aspect of pinnae
Sporangia on the underside of pinnae
General aspect of the fern
Details of the lamina
Entire fronds
Pinnae
Hairs on the stipe and pinnae
Sori
Entire frond
Lateral pinnae
Terminal pinnae
Sori
Entire fronds
Pinnae showing sori on the upper margin
Entire frond
Details of pinnae and sori
General aspect of the fern
Shape of the lamina
Underside of a fertile frond
Details of sori
General aspect of the fern (upper face)
Lower face of a fertile frond
Insertion of a fertile pinnae of the rachis: overview of the false indusium
Sori
Upper face of the frond
Lower face of the frond
Pinnae
Pinnules
General aspect of the frond (upper face)
Lower face of the lamina, showing sori
General aspect of the frond
Base of the lamina showing basal spur
Lower face of the frond
Sori on pinnules
General aspect of the fern (upper face)
General aspect of the fern (lower face)
Lower face of a pinnae showing linear sori
Sori
General aspect of the frond
Lower face of the lamina showing sori
List of descriptors with their discriminating power.
The index value is the ratio between the number of couples with no common values and the total number of couples. For example, the value of the descriptor "habitat" index is 0.12 corresponding to 16 couples with no common values and a total number of 136 available couples.
|
|
1. type of habitat | 16/136 (0.12) |
2. type of terrestrial habitat | 21/120 (0.18) |
3. biotope | 30/136 (0.22) |
4. fronds dimorphism | 52/136 (0.38) |
5. type of dimorphism | 0/6 (0.0) |
6. frond length | 39/136 (0.29) |
7. frond width | 85/136 (0.63) |
8. fronds insertion on the rhizome | 60/136 (0.44) |
9. limb general shape | 102/136 (0.75) |
10. limb length | 56/136 (0.41) |
11. limb width | 82/136 (0.6) |
12. limb length/width ratio | 83/136 (0.61) |
13. limb texture | 72/136 (0.53) |
14. number of limb divisions | 90/136 (0.66) |
15. pinnae length | 38/105 (0.36) |
16. pinnae width | 29/105 (0.28) |
17. pinnae length/width ratio | 44/105 (0.42) |
18. pinnae dimorphism | 36/105 (0.34) |
19. position of the dimorphism | 28/66 (0.42) |
20. basal pinnae basiscopical development | 0/10 (0.0) |
21. terminal pinnae shape | 7/28 (0.25) |
22. pinnae general shape | 38/105 (0.36) |
23. pinnae base shape | 76/105 (0.72) |
24. pinnule general shape | 24/45 (0.53) |
25. pinnae insertion on the rachis | 54/105 (0.51) |
26. pinnae stipe length | 16/28 (0.57) |
27. winged frond | 56/105 (0.53) |
28. frond 3-branched | 10/21 (0.48) |
29. type of veins organization | 96/136 (0.71) |
30. type of dichotomy | 5/15 (0.33) |
31. veins organization | 4/15 (0.27) |
32. limb margin | 83/136 (0.61) |
33. stipe color | 84/136 (0.62) |
34. stipe length | 55/136 (0.4) |
35. rhizome size | 42/136 (0.31) |
36. rhizome habit | 36/136 (0.26) |
37. roots insertion on the rhizome | 0/136 (0.0) |
38. frond type | 0/136 (0.0) |
39. sporangia organization | 30/136 (0.22) |
40. sori position | 14/105 (0.13) |
41. sori position in relation to veins | 0/0 (0.0) |
42. sori position in relation to lamina | 33/91 (0.36) |
43. sori shape | 54/105 (0.51) |
44. indusium | 86/136 (0.63) |
45. indusium opening | 0/1 (0.0) |
46. indusium shape | 0/1 (0.0) |
47. indusium texture | 0/1 (0.0) |
48. covering | 30/136 (0.22) |
49. position of covering | 6/105 (0.06) |
50. rhizome covering | 0/91 (0.0) |
51. rhizome scale’s shape | 48/91 (0.53) |
52. rhizome scale’s color | 32/91 (0.35) |
53. stipe covering | 15/36 (0.42) |
54. stipe scale density | 12/21 (0.57) |
55. stipe muricule size | 0/0 (0.0) |
56. stipe hairs density | 0/0 (0.0) |
57. length of stipe hairs | 0/0 (0.0) |
58. limb covering | 0/6 (0.0) |
59. type of covering | 6/6 (1.0) |
60. limb muricule size | 0/0 (0.0) |
61. farinose coat color | 0/0 (0.0) |
62. limb hairs density | 0/0 (0.0) |
Current nomenclature of the
|
|
||
|
|
|
|
|
|
||
|
|
||
|
|
||
|
|
||
|
|
||
|
|
||
|
|
||
|
|
||
|
|
||
|
|
||
|
- | - | |
|
|
|
|
|
|
||
|
|
||
|
|
|
|
|
|
||
|
|
||
|
|
||
|
|
||
- | - |
|
Number of
Data type: occurences
File: oo_41787.xls