Abstract
Animal pollination is critical for wild plant reproduction and crop yield. However, not all flower visitors are necessarily pollinators. Using cocoa ( Theobroma cacao L.) as a case study, we explored a combination of indicators to assess the effectiveness of flower visitors as pollinators. We first investigated the potential trade-off between the quality and quantity of pollen deposited on the limitation of pollination success in a hand pollination trial under controlled conditions. We found that the fruit set and ovule fecundation rate are mainly driven by the quantity of pollen grain deposited. We then conducted transects and video monitoring in the field in Ecuador to compare the performance of cocoa flower visitors as pollinators using several indicators. We found that small crawling arthropods, in particular ants and aphids, proved to be promising pollinators, with a higher level of pollination with viable pollen being observed on the flowers they visited. Ants were even more efficient than other visitors in depositing significant numbers of pollen grains on the styles. Instead, stingless bees and microdiptera were less efficient when depositing pollen, but complementary to small crawling arthropods in the time of flower visits. Interestingly, we found that others cocoa flower visitors were detrimental to the pollination service, such as hemipteran nymphs and caterpillars. We conclude that cocoa plantation management focused on increasing the complementarity of pollinator visits, acting at different periods of the day and with distinct behaviours, could improve pollination services, and that the use of multiple indicators should be advocated in future research to assess pollinator effectiveness. • Animal pollination is critical in agriculture but not all flowers’ visitors are pollinators. • The success of cocoa pollination is mainly driven by the quantity of pollen grains deposited. • Some cocoa flower visitors are pollinators, but others hinder pollination services. • Ants are more effective than other visitors in depositing pollen grains on the styles. • Cocoa pollination services may rely on the complementarity of several pollinators.
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Claims 8
Claim · model-written summary
Fruit set and ovule fecundation rate in cocoa are mainly driven by the quantity of pollen grains deposited.
- Taxon
- Theobroma cacao
Theobroma cacaoe0 - Treatment
- quantity of pollen grain depositede1
- Outcome
- fruit set and ovule fecundation ratee1
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“cocoa ( Theobroma cacao L.)”
e1“the fruit set and ovule fecundation rate are mainly driven by the quantity of pollen grain deposited”
Claim · model-written summary
Small crawling arthropods, especially ants and aphids, showed a higher level of pollination with viable pollen on the flowers they visited.
- Taxon
- Theobroma cacao
Theobroma cacaoe0 - Interactor
- antsFormicidaee2pollinator
- Second interactor
- aphidsAphidomorphae2pollinator
- Outcome
- pollination with viable pollene2
- Direction
- highere2
- Setting
- in the field in Ecuadore7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“cocoa ( Theobroma cacao L.)”
e2“small crawling arthropods, in particular ants and aphids, proved to be promising pollinators, with a higher level of pollination with viable pollen being observed on the flowers they visited”
e7“in the field in Ecuador”
Claim · model-written summary
Ants deposited more pollen grains on the styles than other cocoa flower visitors.
- Taxon
- Theobroma cacao
Theobroma cacaoe0 - Interactor
- AntsFormicidaee3pollinator
- Outcome
- pollen grains on the stylese3
- Direction
- even more efficiente3
- Comparator
- other visitorse3
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“cocoa ( Theobroma cacao L.)”
e3“Ants were even more efficient than other visitors in depositing significant numbers of pollen grains on the styles”
Claim · model-written summary
Stingless bees and microdiptera were less efficient than ants at depositing pollen in cocoa.
- Taxon
- Theobroma cacao
Theobroma cacaoe0 - Interactor
- stingless beesMeliponinie4pollinator
- Second interactor
- microdipterae4pollinator
- Outcome
- depositing pollene4
- Direction
- less efficiente4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“cocoa ( Theobroma cacao L.)”
e4“stingless bees and microdiptera were less efficient when depositing pollen, but complementary to small crawling arthropods in the time of flower visits”
Claim · model-written summary
Stingless bees and microdiptera were complementary to small crawling arthropods in the time of flower visits.
- Taxon
- Theobroma cacao
Theobroma cacaoe0 - Interactor
- stingless beesMeliponinie4pollinator
- Second interactor
- microdipterae4pollinator
- Outcome
- time of flower visitse4
- Qualifier
- complementary to small crawling arthropodse4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“cocoa ( Theobroma cacao L.)”
e4“stingless bees and microdiptera were less efficient when depositing pollen, but complementary to small crawling arthropods in the time of flower visits”
Claim · model-written summary
Hemipteran nymphs and caterpillars were detrimental to the cocoa pollination service.
- Taxon
- Theobroma cacao
Theobroma cacaoe0 - Interactor
- hemipteran nymphse5other
- Second interactor
- caterpillarse5other
- Outcome
- pollination servicee5
- Direction
- detrimentale5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“cocoa ( Theobroma cacao L.)”
e5“others cocoa flower visitors were detrimental to the pollination service, such as hemipteran nymphs and caterpillars”
Claim · model-written summary
The authors recommend cocoa plantation management that increases the complementarity of pollinator visits to improve pollination services.
- Taxon
- Theobroma cacao
Theobroma cacaoe0 - Treatment
- cocoa plantation management focused on increasing the complementarity of pollinator visitse6
- Outcome
- pollination servicese6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“cocoa ( Theobroma cacao L.)”
e6“cocoa plantation management focused on increasing the complementarity of pollinator visits, acting at different periods of the day and with distinct behaviours, could improve pollination services”
Claim · model-written summary
The authors recommend using multiple indicators to assess pollinator effectiveness in cocoa.
- Taxon
- Theobroma cacao
Theobroma cacaoe0 - Treatment
- the use of multiple indicatorse7
- Outcome
- pollinator effectivenesse7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“cocoa ( Theobroma cacao L.)”
e7“the use of multiple indicators should be advocated in future research to assess pollinator effectiveness”