Research Article |
Corresponding author: George Gardner Brown ( george.brown@embrapa.br ) Academic editor: Fredric Govedich
© 2021 Marcus Vinicius Cremonesi, Alessandra Santos, Danilo Eduardo Rozane, Marie Luise Carolina Bartz, George Gardner Brown.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Cremonesi MV, Santos A, Rozane DE, Carolina Bartz ML, Brown GG (2021) Earthworm species in Musa spp. plantations in Brazil and worldwide. ZooKeys 1033: 1-33. https://doi.org/10.3897/zookeys.1033.54331
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Bananas and plantains are major commodity/food crops that represent an important habitat for earthworms, although so far, no review is available on earthworm communities associated with banana/plantain crops worldwide. The Vale do Ribeira region is among the largest banana producing areas in Brazil, but little is known of the earthworms living there. Hence, the present study assessed earthworm populations and species in three banana plantations and adjacent Atlantic forest fragments along the Ribeira de Iguape River using standard (hand sorting) methodologies. Furthermore, we review earthworm populations reported in banana/plantain plantations worldwide. Only two species (Pontoscolex corethrurus, Amynthas gracilis) belonging to two families (Rhinodrilidae, Megascolecidae) were found in the Ribeira River valley, occurring concurrently. Abundance was low (< 13 indiv. m-2) compared with other banana plantations worldwide, that frequently surpassed 100 indiv. m-2. More than 70 studies reported earthworms from >200 banana plantations in 28 countries, and mean species richness was 2.7 per site, ranging from 1 to 10 species. Exotics predominated in most sites and P. corethrurus was the most prevalent species encountered. Overall, more than 104 species from 10 families were reported, with around 61 native and 43 exotic widespread species, mainly of the Megascolecidae, Lumbricidae and Acanthodrilidae families. Richness was highest in India (27 spp.) and the Canary Islands (25 spp.), but native species dominated only in a few countries and sites, while exotics were prevalent especially in island countries and Brazil. Lower-input practices appear to be important for earthworm communities and banana plantations can have large earthworm populations in some cases, which may be contributing to soil processes and plant production, topics that deserve further attention. However, many important banana-producing countries have not yet been evaluated, so further work is warranted, both in terms of applied ecology and biodiversity.
Annelida, banana, biodiversity, Oligochaeta, plantain, Pontoscolex corethrurus
Bananas and plantains are large, perennial herbs belonging to the genus Musa, that evolved in Indochina and Southeast Asia, but with major secondary diversification in Africa, India and the Caribbean (
Throughout much of LAC, bananas and plantains are still cultivated at the subsistence level, often in agroforestry systems (
Earthworms are essential service providers for terrestrial ecosystems (
The Vale do Ribeira region, located in northeastern Paraná State and southern São Paulo State, has extensive areas (over 36,000 hectares;
Three counties in the lower Ribeira River valley, all of them in the State of São Paulo were selected for this study: Eldorado, Sete Barras and Registro (Fig.
Land use system, watershed number (WN), age of the land use, geographic coordinates and soil types according to FAO classification (
Site | County | System | WN1 | Age (yrs) | Latitude, Longitude | Soil types |
---|---|---|---|---|---|---|
1 | Eldorado | Banana | 344 | 50 | 24°29'35"S, 48°02'10"W | Cambisols |
2 | Eldorado | Atlantic forest | 344 | > 50 | 24°30'09"S, 48°02'30"W | Cambisols |
3 | Sete Barras | Banana | 422 | 15 | 24°23'34"S, 47°53'51"W | Cambisols |
4 | Sete Barras | Atlantic forest | 422 | > 50 | 24°23'30"S, 47°53'22"W | Cambisols |
5 | Registro | Banana | 379 | 40 | 24°26'56"S, 47°49'41"W | Cambisols / Histosols |
6 | Registro | Atlantic forest | 389 | 45 | 24°26'47"S, 47°49'23"W | Cambisols / Histosols |
Earthworms were collected using an adaptation of the standard sampling method proposed by the Tropical Soil Biology and Fertility (TSBF) Programme (
Both the common and scientific names of banana were used for a bibliographic search online using the keywords for bananas and plantains in English, Portuguese, French and Spanish: Musa (genus), Musa acuminata, Musa balbisiana, banana, banane, banano, plátano and plantain. These were then crossed with the common names of earthworms in these languages: earthworms, minhoca, oligochaeta, oligoqueta, vers de terre and lombriz de tierra. Online scientific databases Web of Science, Science Direct, Scielo, google academic and the Base de Dados de Teses e Dissertações (BDTD – Thesis and Dissertation Database) of Brazil were consulted. All the resulting publications were consulted and those containing data on earthworm abundance (density and/or biomass) or species identification were selected and these data extracted, as well as information on sampling sites (counties, countries, management practices of the plantations). Earthworm species were separated into different families and into native or exotic to the region of occurrence, and species richness per site and for each group (native, exotic), when available. Although we treated bananas and plantains separately when possible, for most of the analysis we considered them together, since not all publications provided details regarding the types of bananas cultivated, and even plantains are often called ‘bananas.’ Details on the species and management data obtained and presented in this paper are available for download online from the open access repository Mendeley Data at http://dx.doi.org/10.17632/p8ywsnj8c5.1 (
Quantitative data on the earthworm abundance and biomass obtained from the literature and from the present study were treated as follows. Means of earthworm abundance (no. individuals m-2) and biomass (fresh mass in gm-2) were calculated per sampling site (plantation), using data from the present study. When quantitative data from the literature was available for the individual site, it was used as is. When only means for several plantations in the same general location were provided, these were also used. As the interest of the present study was more at the spatial (site-level) rather than the temporal scale, when samples were taken on multiple occasions, and individual means per sampling date were not available, overall means were used. When taken in wet and dry seasons, both values were used as an interval of abundance and biomass (when measured).
Pontoscolex (Pontoscolex) corethrurus (Müller, 1857)
COFMBRSP0231, 1 individual in Atlantic Forest, HMN 389, Registro – SP (24°26'16.85"S, 47°49'31.71"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0232, 2 individuals in Atlantic Forest, HMN 389, Registro – SP (24°26'16.82"S, 47°49'31.71"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0233, 2 individuals in Atlantic Forest, HMN 389, Registro – SP (24°26'16.28"S, 47°49'32.52"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0235, 2 individuals in Atlantic Forest, HMN 389, Registro – SP (24°26'15.71"S, 47°49'33.32"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0236, 1 individual in Atlantic Forest, HMN 389, Registro – SP (24°26'14.57"S, 47°49'35.35"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0238, 2 individuals in banana field, HMN 379, Registro – SP (24°26'54.25"S, 47°49'38.12"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0239, 1 individual in banana field, HMN 379, Registro – SP (24°26'54.81"S, 47°49'39.41"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0240, 1 individual in Atlantic Forest, HMN 422, Sete Barras – SP (24°23'44.43"S, 47°55'11.56"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0241, 1 individual in Atlantic Forest, HMN 422, Sete Barras – SP (24°23'44.46"S, 47°55'11.49"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0242, 2 individuals in Atlantic Forest, HMN 422, Sete Barras – SP (24°23'43.79"S, 47°55'24.53"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0244, 1 individual in Atlantic Forest, HMN 422, Sete Barras – SP (24°23'43.93"S, 47°55'10.17"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0245, 3 individuals in Atlantic Forest, HMN 422, Sete Barras – SP (24°23'44.33"S, 47°55'09.65"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0248, 1 individual in Atlantic Forest, HMN 422, Sete Barras – SP (24°23'44.90"S, 47°55'08.92"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0249, 1 individual in banana field, HMN 422, Sete Barras – SP (24°23'38.61"S, 47°55'23.49"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0251, 1 individual in banana field, HMN 422, Sete Barras – SP (24°23'43.01"S, 47°55'24.52"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0252, 3 individuals in banana field, HMN 422, Sete Barras – SP (24°23'42.54"S, 47°55'25.32"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0253, 1 individual in Atlantic Forest, HMN 344, Eldorado – SP (24°29'57.34"S, 48°02'41.68"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0255, 1 individual in Atlantic Forest, HMN 344, Eldorado – SP (24°29'55.69"S, 48°02'42.15"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0256, 2 individuals in banana field, HMN 344, Eldorado – SP (24°29'36.89"S, 48°02'09.43"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0258, 2 individuals in banana field, HMN 344, Eldorado – SP (24°29'37.11"S, 48°02'10.84"W), 2019, M. Cremonesi, A. Santos colls.
Rhinodrilidae juveniles. COFMBRSP0246, 1 individual in Atlantic Forest, HMN 422, Sete Barras – SP (24°23'44.33"S, 47°55'09.65"W), 2019, M. Cremonesi, A. Santos colls.
Amynthas gracilis (Kinberg, 1867)
COFMBRSP0237, 1 individual in banana field, HMN 379, Registro – SP (24°26'54.25"S, 47°49'38.22"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0250, 3 individuals in banana field, HMN 422, Sete Barras – SP (24°23'38.61"S, 47°55'23.49"W), 2019, M. Cremonesi, A. Santos colls.
Megascolecidae juveniles. COFMBRSP0234, 1 individual in Atlantic Forest, HMN 389, Registro – SP (24°26'16.28"S, 47°49'32.52"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0243, 1 individual in Atlantic Forest, HMN 422, Sete Barras – SP (24°23'44.06"S, 47°55'10.35"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0247, 1 individual in Atlantic Forest, HMN 422, Sete Barras – SP (24°23'44.33"S, 47°55'09.65"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0254, 1 individual in Atlantic Forest, HMN 344, Eldorado – SP (24°29'56.60"S, 48°02'42.23"W), 2019, M. Cremonesi, A. Santos colls. COFMBRSP0257, 1 individual in banana field, HMN 344, Eldorado – SP (24°29'36.89"S, 48°02'09.43"W), 2019, M. Cremonesi, A. Santos colls.
Only two earthworm species belonging to two families (Rhinodrilidae, Megascolecidae) were found at the six sampling sites in the three counties (Table
Earthworm families, species, and richness in banana plantations and Atlantic Forest remnants, in three counties of the Ribeira de Iguape River valley (Eldorado, Sete Barras, Registro). + means presence and – means absence.
Earthworm family and species | Eldorado | Sete Barras | Registro | |||
---|---|---|---|---|---|---|
Banana | Atlantic Forest | Banana | Atlantic Forest | Banana | Atlantic Forest | |
Megascolecidae | ||||||
Amynthas gracilis | – | – | + | + | + | – |
Megascolecidae juveniles | + | + | – | + | – | + |
Rhinodrilidae | ||||||
Pontoscolex corethrurus | + | + | + | + | + | + |
Rhinodrilidae juveniles | – | – | – | + | – | – |
Species Richness | 2 | 2 | 2 | ≥2 | 2 | 2 |
Most of the individuals collected (76% of the total) were of P. corethrurus, representing 29% of the total abundance in banana crop sites and 46% in Atlantic forest fragments (Fig.
The predominance of P. corethrurus in both native forest and banana plantations of the Ribeira River valley indicate that non-native species have extensively colonized disturbed soils of this region. Nonetheless, this potentially widespread occurrence of exotics should be further evaluated both regionally and nationally, in order to better determine the extent of this phenomenon as well as its possible causes.
Mean overall abundance and biomass of earthworms found in the three sites studied here (6 to 13 indiv. m-2 and 2.5 to 9 g m-2) tended to be quite low compared with others observed overall in Brazil (21 to 459 indiv. m-2 and 3.1 to 177.4 g m-2; see Table
Earthworm abundance and biomass found in banana plantations worldwide, and the predominant species encountered (when available).
Country | Location | Abundance (indiv. m–2) | Biomass (g m–2) | Predominant species | References |
---|---|---|---|---|---|
Brazil | Antonina (Monoculture) | 71 | 35.1 | P. corethrurus |
|
221 | 95.7 | P. corethrurus | |||
86 | 23.8 | P. corethrurus | |||
Antonina (Agroforestry) | 173 | 77.1 | P. corethrurus | ||
338 | 69.6 | P. corethrurus | |||
117 | 43.5 | P. corethrurus | |||
21a | 3.1b | P. corethrurus |
|
||
293a | 34.9b | P. corethrurus | |||
Adrianópolis (Agroforestry) | 211–413c | 37–71.2c | P. corethrurus |
|
|
Barra do Turvo (Agroforestry) | 99–176c | 11.2–17.3c | P. corethrurus | ||
229–459c | 48.3–117.4c | P. corethrurus | |||
Casimiro de Abreu | ~205–440c | – | NA |
|
|
Paraty | 167 | – | NA |
|
|
Eldorado | 8 | 3.9 | P. corethrurus | This study | |
Sete Barras | 13 | 9.0 | P. corethrurus | ||
Registro | 6 | 2.5 | P. corethrurus | ||
Cameroon | Mbalmayo Forest Reserve | 70 | – | Legonodrilus sp. nov. 1, Eminoscolex lamani |
|
121 | – | Legonodrilus sp. nov. 1, Eminoscolex lamani | |||
Campo Ma’an | 16–92d | – | NA |
|
|
Colombia | Quindío (Armenia) | 9–16e | 1.2–3.0e | NAf | Molina and Feijoo (2016) |
Costa Rica | Limón Province (Finca San Pablo) | 83–812g | – | NA |
|
Pueblo Nuevo de Villa Franca de Guácimo, Limón | 29 | 6.2 | NA |
|
|
Cahuita | 350 | 144.6 | P. corethrurus |
|
|
Guadeloupe (France) | Basse-Terre Andosols (mean of 23 sites) | 88 | 23 | NA |
|
Basse Terre Nitisols (mean of 11 sites) | 54 | 17.5 | NA | ||
Capesterre-Belle-Eau (Gloria Bas) | 168 | 27.6 | P. corethrurus |
|
|
Capesterre-Belle-Eau (Source) | 288 | 42.2 | P. corethrurus | ||
Capesterre-Belle-Eau (Bergerie) | 188 | 33.6 | P. corethrurus | ||
Baillif (Sextius) | 336 | 112 | P. corethrurus | ||
Baillif (Grand Canon) | 192 | 70.8 | P. corethrurus | ||
Saint-Claude (Saut d’Eau) | 364 | 46 | P. corethrurus | ||
Ecuador | Latacunga (La Maná) | 168 | – | NA |
|
111 | – | NA | |||
Manabí (El Carmen) | 78 | – | NA | ||
37 | – | NA | |||
El Carmen (Cijádi) | 0–145h | – | NA |
|
|
El Carmen (Nápoles) | 34–144h | – | NA | ||
Santo Domingo de los Tsáchilas (Santa Patrícia) | 83–548h | – | NA | ||
Santo Domingo de los Tsáchilas (La Floresta) | 22–150h | – | NA | ||
India | West Tripura | 16–656i | 4.8–453.6i | P. corethrurus |
|
Rajapalayam | 116 | 48.8 | Lampito mauritii, Perionyx excavatus |
|
|
Ivory Coast | Taabo (Lamto reservation) | 186 | 8.5 | Reginaldia anomala |
|
Martinique (France) | Le Lorrain (Feugère) | 244 | 67.6 | P. corethrurus |
|
Le Lorrain (Bellevue) | 152 | 43.6 | P. corethrurus | ||
Le Lorrain (Limite) | 52 | 26 | P. corethrurus | ||
L’Ajoupa-Bouillon (Allée Domergue 3) | 148 | 49.6 | P. corethrurus | ||
Basse-Pointe (Fromager Rivière) | 80 | 26 | P. corethrurus | ||
Basse-Pointe (Dantu Bas) | 40 | 9 | P. corethrurus | ||
Mexico | Tabasco, Pablo L. Sidar | 25 | 10 | P. corethrurus, Lavellodrilus bonampakensis |
|
Tabasco, Teapa | 116 | 20.8 | P. corethrurus, Drawida barwelli, Polypheretima elongata | ||
117 | 11.8 | Balanteodrilus pearsei, Drawida barwelli | |||
94 | 40.4 | Balanteodrilus pearsei, Polypheretima elongata | |||
125 | 35.6 | P. corethrurus, Drawida barwelli | |||
25 | 8.8 | P. corethrurus, Lavellodrilus bonampakensis |
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||
~350 | 2.5 | Diplotrema murchiei |
|
||
~350 | 9.3 | P. corethrurus | |||
~470 | 16.2 | P. corethrurus | |||
~100 | 11 | P. corethrurus | |||
~80 | 2.8 | P. corethrurus | |||
~125 | 0.8 | Dichogaster sp. | |||
Nicaragua | León (Finca Cony) | 150 | – | NA | Hernández et al. (2015) |
León (Finca San Martín) | 325 | – | NA | ||
León (Finca Santa Isabel) | 50 | – | NA | ||
León (Finca El verdon) | 65 | – | NA | ||
Possoltega (Finca San Joaquin) | 150 | – | NA | ||
Possoltega (Finca Los Ángeles) | 225 | – | NA | ||
Possoltega (Finca Maria de los Ángeles) | 100 | – | NA | ||
Possoltega (Finca Montes Verdes) | 125 | – | NA | ||
Philippines | Davao (Sumitomo Fruits Corporation) | ~85–175j | – | NA |
|
~75–215j | Metaphire cai | ||||
South Africa | Kwazulu-Natal (Eshowe) | 1500k | 180 | Amynthas rodericensis, Amynthas minimus, P. corethrurus |
|
Uganda | Kabanyolo University Farm | 18–207l | 0.1–9.4l | Dichogaster sp. 2, Gordiodrilus sp. 1 |
|
Mabira Forest reserve (1 yr old) | 13 | 0.4 | NA |
|
|
(2 yr old) | 125 | 2.2 | NA | ||
(3 yr old) | 131 | 1.3 | NA | ||
(5 yr old) | 54 | 0.5 | NA | ||
(20 yr old) | 154 | 4.2 | NA |
More than 70 studies were found from 28 countries with data on earthworms in banana and plantain fields (Tables
Earthworm species, richness and number of native and exotic species found in banana plantations under various management practices worldwide.
Country | Location | Management | Culture type | Earthworm species | Richness | Native (N) /Exotic (E) | References |
---|---|---|---|---|---|---|---|
Bangladesh | Lalmonirhat District | NA | NA | Lampito mauritii, Metaphire posthuma, Pontoscolex corethrurus | 3 | 2/1 |
|
Bermuda | Paget Parish | NA | NA | Amynthas rodericensis | 1 | 0/1 |
|
Southampton Parish | NA | NA | Amynthas hupeiensis | 1 | 0/1 |
|
|
Brazil | Antonina, PR | Agroforestry | Monoculture | Amynthas corticis, Pontoscolex corethrurus, two other spp. | 4 | 0/4 |
|
Antonina, PR | Agroforestry | Monoculture | Amynthas gracilis, Pontoscolex corethrurus, two other spp. | 4 | 0/4 |
|
|
Antonina, PR | Agroforestry | Monoculture | Dichogaster spp., Pontoscolex corethrurus | 4 | 0/4 |
|
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Antonina, PR | Agroforestry | Monoculture | Ocnerodrilus occidentalis, Pontoscolex corethrurus, two other spp. | 4 | 0/4 |
|
|
Antonina, PR | Agroforestry | Monoculture | Pontoscolex corethrurus, one other sp. | 2 | 0/2 |
|
|
Antonina, PR | NA | Polyculture | Pontoscolex corethrurus, one other sp. | 2 | 0/2 |
|
|
Antonina, PR | Agroforestry | Polyculture | Dichogaster sp., Pontoscolex corethrurus, and one unidentified sp. | 3 | ?/2 |
|
|
Antonina, PR | Agroforestry | Polyculture | Pontoscolex corethrurus | 1 | 0/1 |
|
|
Adrianópolis, PR | Agroforestry | Polyculture | Amynthas gracilis, Pontoscolex corethrurus | 2 | 0/2 |
|
|
Barra do Turvo, SP | Agroforestry | Polyculture | Amynthas gracilis, Pontoscolex corethrurus | 2 | 0/2 |
|
|
Barra do Turvo, SP | Agroforestry | Polyculture | Amynthas gracilis, Dichogaster sp., Pontoscolex corethrurus | 3 | 0/3 |
|
|
Areia, PB | NA | Polyculture | Amynthas gracilis, Dichogaster affinis, Eudrilus eugeniae, Pontoscolex corethrurus | 4 | 0/4 |
|
|
Eldorado, SP | Conventional | Monoculture | Amynthas gracilis, Pontoscolex corethrurus | 2 | 0/2 | This study | |
Jutaí River margin, AM | NA | NA | Pontoscolex corethrurus | ND | 0/1 |
|
|
Registro, SP | Conventional | Monoculture | Amynthas gracilis, Pontoscolex corethrurus | 2 | 0/2 | This study | |
Sete Barras, SP | Conventional | Monoculture | Amynthas gracilis, Pontoscolex corethrurus | 2 | 0/2 | This study | |
Cameroon | Mbalmayo Forest Reserve (low density cover) | Organic Agroforestry | Monoculture | Dichogaster hauseri, Eminoscolex lamani, Eudrilidae gen. et sp. nov.1 & 2, Legonodrilus sp. nov.1, Malodrilus kamerunensis, Nematogenia panamaensis, Rosadrilus camerunensis | 8 | 7/1 |
|
Mbalmayo Forest Reserve (high density cover) | Organic Agroforestry | Monoculture | Dichogaster annae, Dichogaster bolaui, Dichogaster sp., Eminoscolex lamani, Eudrilidae sp., Eudrilidae gen. et sp. nov. 1, Legonodrilus sp. nov. 1, Nematogenia panamaensis, Ocnerodrilidae gen. et sp. nov., Rosadrilus camerunensis, Scolecillus tantillus | 10 | 7/3 |
|
|
China | Hainan Province | NA | NA | Pheretima montana | ND | 0/1 |
|
Colombia | Quindío, Circasia, Barcelona (La Sofe farm) | NA | Monoculture | Aptodrilus fuhrmanni, Amynthas minimus, Glossodrilus chaguala, Glossodrilus panikita, Martiodrilus quimbayaensis | 5 | 4/1 |
|
Quindío, Circasia, Barcelona (La Sofe farm) | NA | Polyculture | Aptodrilus fuhrmanni, Amynthas minimus, Glossodrilus chaguala, Glossodrilus panikita, Martiodrilus quimbayaensis | 5 | 4/1 |
|
|
Quindío, Circasia, Barcelona (La Sofe farm) | NA | NA | Amynthas gracilis, Periscolex columbianus | 2 | 1/1 |
|
|
Colombia | Armenia, Niagara (La Catalina) | NA | NA | Amynthas gracilis, Glossodrilus griseus, Pontoscolex corethrurus | 3 | 1/2 |
|
Quindio, Calarcá, Quebrada Negra | NA | NA | Glossodrilus griseus | 1 | 1/0 |
|
|
Quindío, Marmato (La Cristalina farm) | NA | Monoculture | Glossodrilus lacteus | 1 | 1/0 |
|
|
Quindío, Marmato (La Cristalina farm) | NA | Polyculture | Glossodrilus lacteus | 1 | 1/0 |
|
|
Quindío, Marmato (La Cristalina farm) | NA | NA | Dichogaster affinis | 1 | 0/1 |
|
|
Armenia, La Revancha (Villa Sofia farm) | NA | NA | Amynthas gracilis, Dichogaster affinis, Dichogaster bolaui, Glossodrilus griseus, Perionyx excavatus | 5 | 1/4 |
|
|
Armenia, La Revancha (Bella Marina farm) | NA | NA | Dichogaster saliens, Periscolex columbianus | 2 | 1/1 |
|
|
Quindío, Armenia, El Rhin | NA | NA | Periscolex columbianus | 1 | 1/0 |
|
|
Quindío, Armenia, La India (La Ermita farm) | NA | NA | Periscolex coreguaje | 1 | 1/0 |
|
|
Circasia, Barcelona Baja rural (Buenos Aires farm) | NA | NA | Amynthas gracilis, Dichogaster saliens, Pontoscolex corethrurus | 3 | 0/3 |
|
|
Quindío, Armenia, La India (La Miranda farm) | NA | NA | Dichogaster saliens | 1 | 0/1 |
|
|
Quindío, Armenia, La Patria | NA | NA | Dichogaster saliens | 1 | 0/1 |
|
|
Costa Rica | Cahuita | NA | NA | Pontoscolex corethrurus | ND | ?/1 |
|
Cuba | Boyeros | Organic | Monoculture | Dichogaster affinis, Dichogaster bolaui, Onychochaeta elegans, Polypheretima elongata, Protozapotecia angelesae | 5 | 2/3 | |
Guadeloupe (France) | Capesterre-Belle-Eau | NA | Monoculture | Pontoscolex corethrurus | ND | ?/1 |
|
Capesterre-Belle-Eau (Gloria Bas) | Conventional | Monoculture | Pontoscolex corethrurus | 1 | 0/1 |
|
|
Capesterre-Belle-Eau (Source) | Conventional | Monoculture | Pontoscolex corethrurus | 1 | 0/1 |
|
|
Capesterre-Belle-Eau (Bergerie) | Agroecological | Monoculture | Pontoscolex corethrurus, unknown sp. 2 | 2 | ? |
|
|
Baillilf (Sextius) | Agroecological | Monoculture | Pontoscolex corethrurus, unknown sp. | 2 | ? |
|
|
Baillilf (Grand Canon) | Agroecological | Monoculture | Pontoscolex corethrurus, unknown sp. 3 | 2 | ? |
|
|
Saint-Claude (Saut d’Eau) | Agroecological | Monoculture | Pontoscolex corethrurus | 1 | 0/1 |
|
|
India | Dakshina Kannada District (Belthangady) | NA | NA | Hoplochaetella kempi | ND | 1/0 |
|
Dakshina Kannada District (Mangalore) | NA | NA | Konkadrilus bahli | ND | 1/0 |
|
|
Dakshina Kannada District (Mangalore) | NA | NA | Dichogaster affinis | ND | 0/1 |
|
|
India | Dakshina Kannada District (Bantwal) | NA | NA | Octochaetona parva | ND | 1/0 |
|
Dakshina Kannada District (sites not detailed) | NA | NA | Amynthas corticis, Hoplochaetella kempi, Hoplochaetella stuarti, Hoplochaetella suctoria, Megascolex konkanensis, Metaphire posthuma, Octochaetona paliensis, Octochaetona parva | ND | 7/1 |
|
|
Kerala (Vellayambalam) | NA | NA | Perionyx excavatus, Pontoscolex corethrurus | 2 | 0/2 |
|
|
Mizoram | NA | Monoculture | Drawida nepalensis, Drawida rangamatiana, Drawida sp., Metaphire houlleti, Perionyx excavatus | 5 | 3/2 |
|
|
Mizoram | NA | Polyculture | Drawida nagana, Drawida sp., Metaphire houlleti, Perionyx excavatus | 4 | 2/2 |
|
|
Rajapalayam | NA | NA | Lampito mauritii, Perionyx excavatus | 2 | 1/1 |
|
|
Udupi District (Adve) | NA | NA | Megascolex konkanensis | 1 | 1/0 |
|
|
Udupi District (Adve) | NA | NA | Metaphire houlleti | 1 | 0/1 |
|
|
Udupi District (Bellibetu) | NA | NA | Metaphire houlleti, Pontoscolex corethrurus | 2 | 0/2 |
|
|
Udupi District (Mudarangadi) | NA | NA | Pontoscolex corethrurus | 1 | 0/1 |
|
|
Udupi District (Nandikur) | NA | NA | Drawida ampullacea, Drawida sulcata, Metaphire peguana | 3 | 3/0 |
|
|
Udupi District (Nandikur) | NA | NA | Drawida ampullacea | 1 | 1/0 |
|
|
Udupi District (Padabettu) | NA | NA | Perionyx excavatus | 1 | 0/1 |
|
|
Udupi District (Yellur) | NA | NA | Mallehulla indica, Megascolex konkanensis | 2 | 2/0 |
|
|
West Tripura (Mohanpur, Maheshkhola, Rastermatha) | Organic | Monoculture | Amynthas alexandri, Drawida assamensis, Drawida papillifer, Eutyphoeus comillahnus, Lampito mauritii, Lennogaster sp., Metaphire houlleti, Metaphire posthuma, Octochaetona beatrix, Perionyx excavatus, Pontoscolex corethrurus | 3–7 | 4/7 |
|
|
Indonesia | Bangkalan (Kamal, Burneh, Socah, Bypass) | NA | NA | Amynthas robustus, Metaphire californica, Metaphire javanica | ND | 1/2 |
|
Bangkalan (Tanah Merah) | NA | NA | Metaphire posthuma | 1 | 0/1 |
|
|
Bangkalan (Labang) | NA | NA | Amynthas robustus, Metaphire javanica, Metaphire californica, Pheretima racemosa | 4 | 2/2 |
|
|
Gresik (Driyorejo, Kedamean, Ngipik, SumengkoLegundi) | NA | NA | Amynthas robustus, Metaphire javanica | ND | 1/1 |
|
|
Gresik (Wringinanamon) | NA | NA | Amynthas robustus, Metaphire javanica, Metaphire posthuma | 3 | 1/2 |
|
|
Sidoarjo (Waru, Taman, Sidoarjo, Tulangan, Tanggulangin, Candi) | NA | NA | Amynthas robustus, Metaphire javanica, Metaphire posthuma | ND | 1/2 |
|
|
Surubaya (Pakal, Benowo, Tandes, Sukolilo, Gubeng, Gununganyar) | NA | NA | Amynthas robustus, Metaphire javanica, Metaphire posthuma | ND | 1/2 |
|
|
Ivory Coast | Lamto region | NA | NA | Dichogaster wenkei, Reginaldia anomala, Stuhlmannia palustris, Stuhlmannia zielae | ND | 4/0 |
|
Jamaica | Clarendon, Crofts Mountain | NA | NA | Drawida barwelli, Polypheretima elongata | 2 | 0/2 |
|
Madagascar | Ambatosoratra Ambatondrazaka | NA | NA | Kynotus sihanakus, Kynotus sp.2 | 2 | 2/0 | Razafindrakoto et al. (2016), |
Malaysia | Serdang, Sengalor (Universiti Putra Malaysia) | NA | NA | Pontoscolex corethrurus | ND | 0/1 | Teng et al. (2006) |
Martinique (France) | Le Lorrain (Feugère) | Conventional | Monoculture | Pontoscolex corethrurus | 1 | 0/1 |
|
Le Lorrain (Limite) | Agroecological | Monoculture | Pontoscolex corethrurus | 1 | 0/1 |
|
|
Le Lorrain (Bellevue) | Conventional | Monoculture | Pontoscolex corethrurus | 1 | 0/1 |
|
|
L’Ajoupa-Bouillon (Allée Domergue 3) | Agroecological | Monoculture | Pontoscolex corethrurus | 1 | 0/1 |
|
|
Basse-Pointe (Fromager Rivière) | Conventional | Monoculture | Pontoscolex corethrurus | 1 | 0/1 |
|
|
Basse-Pointe (Dantu Bas) | Agroecological | Monoculture | Pontoscolex corethrurus | 1 | 0/1 |
|
|
Mexico | Tabasco | NA | Monoculture | Lavellodrilus bonampakensis, Pontoscolex corethrurus | 2 | 1/1 |
|
Tabasco, Teapa B1 | NA | Monoculture | Balanteodrilus pearsei, Drawida barwelli, Polypheretima elongata, Pontoscolex corethrurus, Pontoscolex sp. | 5 | 1/4 | ||
Tabasco, Teapa B2 | NA | Monoculture | Balanteodrilus pearsei, Dichogaster bolaui, Drawida barwelli, Periscolex brachycystis, Polypheretima elongata, Pontoscolex sp. | 6 | 2/4 | ||
Tabasco, Teapa AF1 | Agroforestry | Polyculture | Balanteodrilus pearsei, Dichogaster bolaui, Drawida barwelli, Polypheretima elongata, Pontoscolex corethrurus, Pontoscolex sp. | 6 | 2/4 | ||
Tabasco, Teapa AF2 | Agroforestry | Polyculture | Balanteodrilus pearsei, Dichogaster bolaui, Drawida barwelli, Polypheretima elongata, Pontoscolex corethrurus | 5 | 1/4 | ||
Tabasco, Teapa (site 1) | Conventional | NA | Dichogaster saliens, Diplotrema murchiei, Pontoscolex corethrurus | 3 | ½ |
|
|
Tabasco, Teapa (site 2) | Conventional | NA | Dichogaster saliens, Pontoscolex corethrurus | 2 | 0/2 |
|
|
Tabasco, Teapa (site 3) | Conventional | NA | Diplotrema murchiei, Polypheretima elongata, Pontoscolex corethrurus | 3 | 1/2 |
|
|
Tabasco, Teapa (site 4) | Conventional | Polyculture | Amynthas gracilis, Pontoscolex corethrurus | 2 | 0/2 |
|
|
Tabasco, Teapa (site 5) | Conventional | Polyculture | Dichogaster saliens, Polypheretima elongata, Pontoscolex corethrurus | 3 | 0/3 |
|
|
Tabasco, Teapa (site 6) | Conventional | NA | Dichogaster saliens, Pontoscolex corethrurus | 2 | 0/2 |
|
|
Tabasco, Pablo L. Sidar | NA | Monoculture | Lavellodrilus bonampakensis, Pontoscolex corethrurus | 2 | 1/1 |
|
|
Tamaulipas (Biosphere Reserve “El Cielo”) | NA | NA | Amynthas gracilis | ND | 0/1 |
|
|
Actopan, Ejido Buenavista | NA | NA | Balanteodrilus psammophilus | ND | 1/0 |
|
|
Nicaragua | Managua | NA | NA | Dichogaster bolaui, Periscolex brachycystis | 2 | 1/1 |
|
Peru | Sarita Colonia | NA | Monoculture | Pontoscolex corethrurus and two native spp. | 3 | 2/1 |
|
Philippines | Davao (Sumitomo Fruits Corporation, 15% site) | Conventional | Monoculture | Metaphire sp., Pithemera bicincta, Pontoscolex corethrurus | 3 | 1/2 |
|
Davao (Sumitomo Fruits Corporation, 25% site) | Conventional | Monoculture | Metaphire cai, Metapheretima sp., Perionyx excavatus | 3 | 2/1 |
|
|
Portugal | Madeira Island (Ribeira Brava) | NA | NA | Aporrectodea moebii, Eisenia eisensi, Metaphire californica | 3 | 0/3 |
|
Madeira Island (Funchal) | NA | NA | Amynthas gracilis, Metaphire californica, Ocnerodrilus occidentalis | 3 | 0/3 |
|
|
Portugal | Madeira Island (Santa Cruz) | NA | NA | Amynthas gracilis | 1 | 0/1 |
|
Madeira Island (Terceira Lombada) | NA | NA | Aporrectodea moebii, Eiseniella tetraedra | 2 | 0/2 |
|
|
Madeira Island (Porto Moniz) | NA | NA | Amynthas gracilis, Aporrectodea rosea, Aporrectodea trapezoides, Dendrobaena pseudohortensis | 4 | 0/4 |
|
|
Madeira Island (Terceira Lombada) | NA | NA | Aporrectodea caliginosa, Aporrectodea rosea, Eiseniella tetraedra | 3 | 0/3 |
|
|
Seychelles | Cousine Island | NA | Monoculture | Pontoscolex corethrurus | ND | 0/1 |
|
South Africa | KwaZulu-Natal (Fairfield Farm) | NA | Monoculture | Pontoscolex corethrurus | ND | 0/1 |
|
KwaZulu-Natal (Benhurst Farm) | NA | Monoculture | Pontoscolex corethrurus | ND | 0/1 |
|
|
KwaZulu-Natal (6 sites in Eshowe) | NA | Monoculture | Amynthas corticis, Amynthas minimus, Amynthas rodericensis, Dichogaster bolaui, Pontoscolex corethrurus, and one other sp. | ND | 0/5 |
|
|
Spain | Gomera Island (Agulo) | NA | NA | Amynthas rodericensis, Allolobophora chlorotica, Eiseniella tetraedra, Ocnerodrilus occidentalis | 4 | 0/4 |
|
Gomera Island (Barranco de la Villa) | NA | NA | Bimastos rubidus, Ocnerodrilus occidentalis, Pithemera bicincta | 3 | 0/3 |
|
|
Gomera Island (Barranco del Valle) | NA | NA | Allolobophora chlorotica, Metaphire californica | 2 | 0/2 |
|
|
Gomera Island (Casas de Aluce) | NA | NA | Aporrectodea rosea, Microscolex phosphoreus | 2 | 0/2 |
|
|
Gomera Island (Cabo Verde) | NA | NA | Amynthas gracilis, Bimastos rubidus | 2 | 0/2 |
|
|
Gomera Island (Costa Agulo) | NA | NA | Aporrectodea trapezoides, Amynthas rodericensis, Bimastos rubidus, Ocnerodrilus occidentalis | 4 | 0/4 |
|
|
Gomera Island (El Molinito) | NA | NA | Amynthas morrisi, Microscolex phosphoreus | 2 | 0/2 |
|
|
Gomera Island (Hermigua) | NA | NA | Aporrectodea rosea, Bimastos rubidus, Eisenia fetida, Ocnerodrilus occidentalis | 4 | 0/4 |
|
|
Gomera Island (Laguna de Santiago) | NA | NA | Amynthas morrisi, Aporrectodea rosea, Aporrectodea trapezoides, Bimastos rubidus, Dendrobaena hortensis, Dichogaster affinis, Metaphire californica, Pithemera bicincta | 9 | 0/9 |
|
|
Gomera Island (Playa de Santiago) | NA | NA | Ocnerodrilus occidentalis | 1 | 0/1 |
|
|
Gomera Island (Seimal) | NA | NA | Eiseniella tetraedra, Metaphire californica, Microscolex phosphoreus | 3 | 0/3 |
|
|
Gomera Island (Taguluche) | NA | NA | Amynthas morrisi, Allolobophora chlorotica, Octalasion lacteum | 3 | 0/3 |
|
|
Gomera Island (Valle Gran Rey) | NA | NA | Allolobophora chlorotica, Aporrectodea trapezoides, Dendrobaena hortensis, Eisenia fetida, Microscolex dubius, Pithemera bicincta | 5 | 0/5 |
|
|
Gran Canaria (Lomo del Galeón) | NA | NA | Ocnerodrilus occidentalis | 1 | 0/1 |
|
|
Gran Canaria (Los Llanos) | NA | NA | Ocnerodrilus occidentalis, Pithemera bicincta | 2 | 0/2 |
|
|
Gran Canaria Island (Bañaderos) | NA | NA | Metaphire californica | 1 | 0/1 |
|
|
Gran Canaria Island (Barranco Guiniguada) | NA | NA | Amynthas morrisi | 1 | 0/1 |
|
|
Gran Canaria Island (Frontón) | NA | NA | Amynthas gracilis | 1 | 0/1 |
|
|
Gran Canaria Island (Galdar) | NA | NA | Amynthas morrisi | 1 | 0/1 |
|
|
Spain | Gran Canaria Island (Hoya Mondondo) | NA | NA | Pithemera bicincta | 1 | 0/1 |
|
Gran Canaria Island (La Aldea) | NA | NA | Dichogaster affinis | 1 | 0/1 |
|
|
Gran Canaria Island (Pedrazo) | NA | NA | Pithemera bicincta | 1 | 0/1 |
|
|
Gran Canaria Island (Tenoya) | NA | NA | Amynthas morrisi | 1 | 0/1 |
|
|
Hierro Island (Los Mocanes) | NA | NA | Ocnerodrilus occidentalis | 1 | 0/1 |
|
|
Hierra Island (NE tip) | NA | NA | Microscolex phosphoreus | ND | 0/1 |
|
|
La Palma Island (Barranco de las Angustias) | NA | NA | Amynthas gracilis | 1 | 0/1 |
|
|
La Palma Island (Barranco Nogales) | NA | NA | Amynthas gracilis | 1 | 0/1 |
|
|
La Palma Island (El Socorro) | NA | NA | Pithemera bicincta | 1 | 0/1 |
|
|
La Palma Island (La Caldereta) | NA | NA | Amynthas morrisi, Metaphire californica | 2 | 0/2 |
|
|
La Palma Island (Los Cancajos) | NA | NA | Amynthas morrisi | 1 | 0/1 |
|
|
La Palma Island (Los Llanos de Aridane) | NA | NA | Amynthas morrisi, Metaphire californica | 2 | 0/2 |
|
|
La Palma Island (Tazacorte) | NA | NA | Amynthas gracilis, Amynthas morrisi, Metaphire californica | 3 | 0/3 |
|
|
Tenerife Island (Abama) | NA | Monoculture | Aporrectodea rosea, Dendrobaena hortensis, Eisenia andrei, Microscolex dubius | 4 | 0/4 |
|
|
Tenerife Island (Adeje) | NA | NA | Ocnerodrilus occidentalis | 1 | 0/1 |
|
|
Tenerife Island (Bajamar) | NA | Monoculture | Amynthas morrisi, Aporrectodea rosea, Dichogaster affinis, Eisenia andrei, Microscolex phosphoreus, Ocnerodrilus occidentalis | 6 | 0/6 |
|
|
Tenerife Island (Barranco de Santos) | NA | Monoculture | Amynthas morrisi, Aporrectodea rosea, Bimastos rubidus, Eisenia andrei, Microscolex dubius, Pithemera bicincta | 6 | 0/6 |
|
|
Tenerife Island (Barranco del Inglés) | NA | Monoculture | Aporrectodea rosea, Aporrectodea trapezoides, Eisenia andrei, Microscolex dubius | 4 | 0/4 |
|
|
Tenerife Island (Barranco la Atalaya) | NA | Monoculture | Aporrectodea rosea, Pithemera bicincta | 2 | 0/2 |
|
|
Tenerife Island (Barranco las Galletas) | NA | Monoculture | Aporrectodea rosea, Eisenia andrei, Ocnerodrilus occidentalis | 3 | 0/3 |
|
|
Tenerife Island (Barranco San Felipe) | NA | Monoculture | Amynthas gracilis, Eisenia andrei, Pithemera bicincta | 3 | 0/3 |
|
|
Tenerife Island (Buenavista del Norte) | NA | NA | Ocnerodrilus occidentalis | 1 | 0/1 |
|
|
Tenerife Island (Casablanca) | NA | Monoculture | Amynthas corticis, Aporrectodea rosea, Eisenia andrei, Ocnerodrilus occidentalis | 4 | 0/4 |
|
|
Tenerife Island (Costa Valle Guerra) | NA | Monoculture | Amynthas gracilis | 1 | 0/1 |
|
|
Tenerife Island (El Puente) | NA | Monoculture | Amynthas gracilis, Aporrectodea rosea, Eisenia andrei, Microscolex phosphoreus, Ocnerodrilus occidentalis | 5 | 0/5 |
|
|
Tenerife Island (El Rincón) | NA | Monoculture | Amynthas gracilis, Bimastos rubidus, Dendrobaena cognetti, Microscolex dubius, Microscolex phosphoreus, Octodrilus complanatus | 6 | 0/6 |
|
|
Spain | Tenerife Island (El Socorro) | NA | NA | Pithemera bicincta | 1 | 0/1 |
|
Tenerife Island (Fañabé) | NA | Monoculture | Amynthas corticis, Aporrectodea rosea, Dichogaster affinis, Eisenia andrei, Ocnerodrilus occidentalis | 5 | 0/5 |
|
|
Tenerife Island (Güimar) | NA | NA | Dichogaster affinis, Ocnerodrilus occidentalis | 2 | 0/2 |
|
|
Tenerife Island (Iboybo) | NA | Monoculture | Aporrectodea rosea, Eisenia andrei, Ocnerodrilus occidentalis | 3 | 0/3 |
|
|
Tenerife Island (Icod de Los Vinos) | NA | Monoculture | Dendrobaena cognetti, Bimastos rubidus, Octodrilus complanatus, Ocnerodrilus occidentalis | 4 | 0/4 |
|
|
Tenerife Island (Igueste) | NA | Monoculture | Allolobophora chlorotica, Aporrectodea rosea, Aporrectodea trapezoides, Pontoscolex corethrurus, Ocnerodrilus occidentalis | 5 | 0/5 |
|
|
Tenerife Island (La Hondura) | NA | Monoculture | Amynthas morrisi | 1 | 0/1 |
|
|
Tenerife Island (La Longuera) | NA | Monoculture | Amynthas morrisi, Aporrectodea rosea, Eisenia fetida, Microscolex dubius, Octodrilus complanatus | 5 | 0/5 |
|
|
Tenerife Island (La Matanza) | NA | Monoculture | Bimastos rubidus, Eisenia andrei, Microscolex phosphoreus | 3 | 0/3 |
|
|
Tenerife Island (La Montañeta) | NA | NA | Pithemera bicincta | 1 | 0/1 |
|
|
Tenerife Island (La Vera) | NA | Monoculture | Bimastos rubidus, Eisenia andrei, Microscolex phosphoreus | 3 | 0/3 |
|
|
Tenerife Island (Las Arenas) | NA | NA | Amynthas morrisi | 1 | 0/1 |
|
|
Tenerife Island (Las Galletas) | NA | Monoculture | Eisenia andrei, Bimastos eiseni, Ocnerodrilus occidentalis | 3 | 0/3 |
|
|
Tenerife Island (Las Madrigueras) | NA | NA | Amynthas morrisi | 1 | 0/1 |
|
|
Tenerife Island (Los Quintos) | NA | Monoculture | Dendrobaena cognetti, Bimastos rubidus, Microscolex phosphoreus, Pithemera bicincta, Ocnerodrilus occidentalis | 5 | 0/5 |
|
|
Tenerife Island (Los Realejos) | NA | NA | Pithemera bicincta | 1 | 0/1 |
|
|
Tenerife Island (Los Rechazos) | NA | Monoculture | Aporrectodea trapezoides, Bimastos rubidus, Eisenia fetida, Octodrilus complanatus, Pithemera bicincta | 5 | 0/5 |
|
|
Tenerife Island (Los Silos) | NA | Monoculture | Amynthas morrisi, Aporrectodea rosea, Dichogaster affinis, Eisenia andrei, Ocnerodrilus occidentalis | 5 | 0/5 |
|
|
Tenerife Island (Loss Llanos) | NA | Monoculture | Amynthas morrisi, Bimastos rubidus, Eisenia andrei, Pithemera bicincta | 4 | 0/4 |
|
|
Tenerife Island (Playa de las Aguas) | NA | Monoculture | Amynthas morrisi, Eisenia andrei, Pithemera bicincta | 3 | 0/3 |
|
|
Tenerife Island (Playa de San Juan) | NA | Monoculture | Aporrectodea rosea, Dendrobaena hortensis, Bimastos rubidus, Eisenia andrei | 4 | 0/4 |
|
|
Tenerife Island (Playa San Marcos) | NA | Monoculture | Pithemera bicincta, Bimastos rubidus, Microscolex phosphoreus | 3 | 0/3 |
|
|
Tenerife Island (Puertito de Gilimar) | NA | Monoculture | Microscolex phosphoreus, Pithemera bicincta, Ocnerodrilus occidentalis | 3 | 0/3 |
|
|
Tenerife Island (Puerto de Santiago) | NA | Monoculture | Amynthas morrisi | 1 | 0/1 |
|
|
Tenerife Island (Punta del Hidalgo) | NA | Monoculture | Amynthas gracilis, Ocnerodrilus occidentalis | 2 | 0/2 |
|
|
Tenerife Island (San Andrés) | NA | Monoculture | Amynthas morrisi, Aporrectodea rosea, Microscolex phosphoreus, Ocnerodrilus occidentalis | 4 | 0/4 |
|
|
Tenerife Island (San Bernardo) | NA | Monoculture | Amynthas corticis, Amynthas morrisi, Aporrectodea rosea, Eisenia andrei | 4 | 0/4 |
|
|
Spain | Tenerife Island (San Juan de la Rambla) | NA | Monoculture | Amynthas gracilis, Bimastos rubidus, Dendrobaena hortensis, Eisenia fetida, Pithemera bicincta | 5 | 0/5 |
|
Tenerife Island (San Pedro de Daute) | NA | Monoculture | Amynthas morrisi, Aporrectodea rosea, Ocnerodrilus occidentalis | 3 | 0/3 |
|
|
Tenerife Island (Santo Domingo) | NA | Monoculture | Dendrobaena cognetti, Microscolex dubius, Microscolex phosphoreus | 3 | 0/3 |
|
|
Tenerife Island (Taganana) | NA | Monoculture | Amynthas morrisi | 1 | 0/1 |
|
|
Tenerife Island (Tejina) | NA | Monoculture | Amynthas corticis, Pithemera bicincta | 2 | 0/2 |
|
|
Taiwan | Central region | NA | NA | Pontoscolex corethrurus | ND | 0/1 |
|
Uganda | Kabanyolo University Farm | NA | NA | Dichogaster sp. 1, Dichogaster sp. 2, Gordiodrilus sp., Pygmaeodrilus sp., Polytoreutus sp. 1 | 5 | 5/0 |
|
Overall, ≥104 earthworm species from 10 earthworm families were recorded from banana/plantain fields worldwide, of which around 61 (59%) were native and 43 exotic to the sampling sites (Table
High species richness was also detected overall in Spain (25), mainly due to the higher sampling effort involving a large number of sites in the Canary Islands. However, all of the species encountered on the islands offshore of Africa were exotic, their introduction having been stimulated over centuries of human colonization bringing in exotic soils and crops (
Number of quantitative (with abundance data) and qualitative (where species were identified) sampling sites and earthworm species (total, native, and exotic) and families found in banana plantations in different countries of the world.
Country | No. sites: Quant./Qual.1 | Total No. species | Native | Exotic | Families |
---|---|---|---|---|---|
Asia | 6/≥47 | 35 | 22 | 13 | 5 |
Bangladesh | 0/1 | 3 | 1 | 2 | 2 |
China | 0/1 | 1 | 0 | 1 | 1 |
India | 4/≥20 | 27 | 20 | 7 | 5 |
Indonesia | 0/23 | 5 | 1 | 4 | 1 |
Malaysia | 0/1 | 1 | 0 | 1 | 1 |
Philippines | 2/2 | 6 | 3 | 3 | 2 |
Taiwan | 0/1 | 1 | 0 | 1 | 1 |
Africa | 33/97 | 50 | 20 | 30 | 7 |
Cameroon | 10/2 | 12 | 9 | 3 | 3 |
Canary Islands (Spain)2 | |||||
Gomera | 0/13 | 18 | 0 | 18 | 4 |
Gran Canaria | 0/10 | 6 | 0 | 6 | 3 |
Hierro | 0/2 | 2 | 0 | 2 | 2 |
La Palma | 0/7 | 3 | 0 | 3 | 1 |
Tenerife | 0/45 | 19 | 0 | 19 | 5 |
Ivory Coast | 1/1 | 4 | 4 | 0 | 3 |
Madagascar | 0/1 | 2 | 2 | 0 | 1 |
Madeira (Portugal)2 | 0/6 | 10 | 0 | 10 | 3 |
Seychelles | 0/1 | 1 | 0 | 1 | 1 |
South Africa | 6/8 | 5 | 0 | 5 | 3 |
Uganda | 6/1 | 5 | 5 | 0 | 3 |
North America | 12/16 | 14 | 7 | 7 | 4 |
Bermuda | 0/2 | 2 | 0 | 2 | 1 |
Mexico | 12/14 | 12 | 5 | 7 | 4 |
Central America/Caribbean | 53/≥17 | ≥10 | 4 | ≥6 | 4 |
Costa Rica | ≥5/1 | 1 | 0 | 1 | 1 |
Cuba | 1/1 | 5 | 2 | 3 | 3 |
Dominica | 1/0 | 2 | 1? | 1 | 2 |
Guadeloupe (France) | 40/7 | 4? | ? | ≥1 | ≥1 |
Martinique (France) | 6/6 | 1 | 0 | 1 | 1 |
Jamaica | 0/1 | 2 | 0 | 2 | 2 |
Nicaragua | 0/1 | 2 | 1 | 1 | 2 |
South America | 49/33 | 20 | 10 | 10 | 6 |
Brazil | 16/16 | 7 | 0 | 7 | 5 |
Colombia | 32/15 | 15 | 8 | 7 | 4 |
Peru | 1/1 | 3 | 2 | 1 | ≥1 |
Total | 153/210 | ≥104 | ≥61 | ≥43 | 10 |
Species richness in individual banana/plantain fields was measured in 166 of the 210 sites, and was generally very low, with an overall mean of 2.7 species per site worldwide, of which less than one (0.5) was native and 2.1 were exotic (full dataset in
There was a clear positive relationship between the number of sites sampled in each country and the total number of species encountered (r = 0.7, p< 0.01), particularly for exotic (r = 0.78, p < 0.01) species (Fig.
A Relationship between species richness (total, native, and exotic species) and the number of sampling sites in each world country (data from Table
Of the over 100 species found in banana and plantain fields worldwide, most belonged to the Megascolecidae (22%), Lumbricidae (17%) and Acanthodrilidae (16%) families (
The most commonly encountered earthworm species in banana plantations was P. corethrurus (11%), found in 15 countries, mainly in Latin America, but also in places as far away as South Africa, India, Bangladesh, Malaysia, Philippines and Taiwan. Interestingly, 37 out of 54 sites (69%) that identified earthworm species reported P. corethrurus as dominant in the banana plantations (Table
Although P. corethrurus may affect soil physical properties negatively by increasing soil compaction under some conditions, it can also positively affect biogeochemical processes, microbial activity, plant production, and soil recovery (see review in
A total of 31 studies performed in 153 sites and 15 countries (Table
Overall earthworm abundance ranged from a minimum of 0 (
These high earthworm abundances and biomasses may be contributing significantly to soil processes (bioturbation, nutrient cycling) in banana/plantain fields, as biomasses over 17 g m-2 and above 32 g m-2 are known to lead to moderate (20–40%) and important (>40%) grain production increases, respectively (
Finally, 18 of the major banana-producing countries in the world (34 countries with >30,000 ha in production, or >1 Million T bananas produced yr-1;
Earthworms are an important component of banana and plantain fields worldwide and deserve further attention by taxonomists, ecologists and agronomists. Under some conditions, especially in lower-input polycultures, their abundance and biomass may reach high values and contribute significantly to soil processes and plant production. More than 70 studies performed in over 200 banana plantations of 28 countries found >100 species (around 60% of them native) from 10 families, although species richness in each sited tended to be low (generally <3 species) and exotic species predominated (particularly P. corethrurus). However, as many important banana-producing countries have not yet been evaluated, further work is warranted in order to better understand the earthworm communities and their functional roles in plantain/banana fields, and the role of management practices in affecting their populations and diversity worldwide.
This study was part of the PhD of the first author at the Federal University of Paraná, where both he and AS received CAPES scholarships. We also gratefully acknowledge financial support from the Fundo Estadual de Recursos Hídricos (Proc. FEHIDRO 102/2016), the CNPq for Profix support (441930/2020-4) and a fellowship to GGB (310690/2017-0), Embrapa Forestry and UFPR for laboratorial and logistical support, and the farmers for permission to sample on their properties.