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Spatiotemporal pattern of specialization of sunbird-plant networks on Mt. Cameroon

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Differences in interaction specializations between nectarivorous birds and plants across continents serve as common examples of evolutionary trajectory specificity. While New World hummingbird-plant networks have been extensively studied and are considered highly specialized, knowledge on the network specialization of their Old World counterparts, sunbirds (Nectariniidae), remains limited. A few studies from tropical Africa indicate that sunbird-plant networks are rather generalized. Unfortunately, these studies are limited to dry seasons and high elevations at the tree line, environments where niche-based hypotheses also often predict lower resource partitioning. In our study, we explored the specialization of sunbird-plant networks and their spatiotemporal variability on Mt. Cameroon (Cameroon). Using a combination of automatic video recordings and personal observations, we constructed eight comprehensive sunbird-plant networks in four forest types at different elevations in both the dry and wet seasons. As reported in previous studies, the montane forest plants, birds and whole networks were highly generalized. Nevertheless, we observed a much higher specialization in forests at lower elevations. Except at the lowest altitude, the wet season was also characterized by higher specialization. While less specialized flowering trees dominated in the dry season networks, more specialized herbs and shrubs were visited by birds during the wet season. As our findings do not support the generally accepted assumption that Old World bird-plant networks are rather generalized, we need further studies to understand the differences in bird-plant specializations on individual continents.
Sunbird-plant networks on Mt. Cameroon at four elevations in dry and wet seasons. LF – lowland forest; MEF – mid-elevation forest; SF – submontane forest; MF – montane forest. PLANTS: Aca dec—Acanthopale decempedalis; Afr sp.—Aframomum sp.; Ant sca—Anthocleista scandens; Ant fra—Anthonotha fragrans; Ber bra—Berlinia bracteosa; Ber rac—Bertiera racemosa; Bri owa—Brillantaisia owariensis; Cha lai—Chassalia laikomensis; Cla ani—Clausena anisata; Cle syl—Clerodendrum sylvanum; Cos dub—Costus dubius; Cor aur—Cordia aurantiaca; Dic ves—Dicranolepis vestita; Dis gra—Dischistocalyx grandiflora; Eng gab—Englerina gabonensis; Hei cri—Heinsia crinita; Hug mic—Hugonia micans; Hyp tri—Hypoestes triflora; Imp bur—Impatiens burtonii; Imp hia—Impatiens hians var. hians; Imp eti—Impatiens etindensis; Imp fri—Impatiens frithii; Imp nia—Impatiens niamniamensis; Imp sak—Impatiens sakeriana; Ixo fol—Ixora foliosa; Ixo gui—Ixora guineensis; Jas pre—Jasminum preussii; Kig afr—Kigelia africana; Lee gui—Leea guineensis; Mim sol—Mimulopsis solmsii; Mus ten—Mussaenda tenuiflora; Nux con—Nuxia congesta; Pav rig—Pavetta rigida; Phr kam—Phragmanthera kamerunensis; Ple dec—Plectranthus decurrens; Ple kam—Plectranthus kamerunensis; Psy hyp—Psychotria hypsophila; Psy dun—Psydrax dunlapii; Rhi sp.—Rhipidoglossum sp.; Sab pil—Sabicea pilosa; Sch mag—Schumanniophyton magnificum; Sch aby—Schefflera abyssinica; Syz sta—Syzygium staudtii; Syz sp.—Syzygium sp.; Tab bra—Tabernaemontana brachyantha; Tab ven—Tabernaemontana ventricosa; Tho san—Thonningia sanguinea; Thu fas—Thunbergia fasciculata; Voa afr—Voacanga africana; SUNBIRDS: Ant col—Anthodiaeta collaris; Ant rec – Anthreptes rectirostris; Ant sei—Anthreptes seimundi; Cha rub—Chalcomitra rubescens; Cin bat—Cinnyris batesi; Cin joh—Cinnyris johannae; Cin min—Cinnyris minullus; Cin urs—Cinnyris ursulae; Cin rei—Cinnyris reichenowi; Cya cya—Cyanomitra cyanolaema; Cya oli—Cyanomitra olivacea; Cya ori—Cyanomitra oritis
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Oecologia (2022) 199:885–896
https://doi.org/10.1007/s00442-022-05234-4
PLANT-MICROBE-ANIMAL INTERACTIONS – ORIGINAL RESEARCH
Spatiotemporal pattern ofspecialization ofsunbird‑plant networks
onMt. Cameroon
ŠtěpánJaneček1 · KryštofChmel1· JiříMlíkovský1· GuillermoUceda‑Gómez1· PetraJanečková1·
NestoralTajaochaFominka1,2· MarcusMokakeNjie1,3· FrancisLumaEwome1
Received: 11 August 2021 / Accepted: 26 July 2022 / Published online: 10 August 2022
© The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2022
Abstract
Differences in interaction specializations between nectarivorous birds and plants across continents serve as common examples
of evolutionary trajectory specificity. While New World hummingbird-plant networks have been extensively studied and
are considered highly specialized, knowledge on the network specialization of their Old World counterparts, sunbirds (Nec-
tariniidae), remains limited. A few studies from tropical Africa indicate that sunbird-plant networks are rather generalized.
Unfortunately, these studies are limited to dry seasons and high elevations at the tree line, environments where niche-based
hypotheses also often predict lower resource partitioning. In our study, we explored the specialization of sunbird-plant net-
works and their spatiotemporal variability on Mt. Cameroon (Cameroon). Using a combination of automatic video recordings
and personal observations, we constructed eight comprehensive sunbird-plant networks in four forest types at different eleva-
tions in both the dry and wet seasons. As reported in previous studies, the montane forest plants, birds and whole networks
were highly generalized. Nevertheless, we observed a much higher specialization in forests at lower elevations. Except at
the lowest altitude, the wet season was also characterized by higher specialization. While less specialized flowering trees
dominated in the dry season networks, more specialized herbs and shrubs were visited by birds during the wet season. As
our findings do not support the generally accepted assumption that Old World bird-plant networks are rather generalized, we
need further studies to understand the differences in bird-plant specializations on individual continents.
Keywords Bird pollination· Ecological network· Elevation· Sunbird· Seasonality
Introduction
Specialization of plant-pollinator networks is a fundamental
ecosystem characteristic. Knowledge on this specialization
and how it is affected by various environmental conditions is
significant from many points of view. It is important not only
for conservation purposes (e.g., for estimating the extinction
risk of individual organisms, Vanbergen etal. 2017) and for
predicting the effects of future climate changes on plant-
pollinator communities (Hoiss etal. 2015) but also for a gen-
eral understanding of evolutionary and speciation processes
(Ramírez etal. 2011).
Specialization in plant-pollinator networks reflects
resource (niche) partitioning among species, which can be
driven by the tendency of plants and pollinators to use only
a subset of potential resources and by interplant or inter-
pollinator interactions such as competition (Blüthgen etal.
2006). Some of the most studied networks are those among
plants and nectarivorous birds. Nevertheless, whereas
researchers have often targeted hummingbird-plant interac-
tions, networks including other groups of nectarivorous birds
remain underexplored. A few studies on African sunbird-
plant networks indicate that they are less specialized than
hummingbird-plant networks (Zanata etal. 2017; Nsor etal.
2019). Nevertheless, other studies revealed many similarities
that should result in a similar degree of specialization. For
Communicated by David M Watson .
* Štěpán Janeček
stepan.janecek@natur.cuni.cz
1 Department ofEcology, Faculty ofScience, Charles
University, Viničná 7, 12844Prague2, CzechRepublic
2 Department ofZoology andAnimal Physiology, Faculty
ofScience, University ofBuea, P.O. Box63, Buea,
Cameroon
3 National Forestry School, Mbalmayo, P.O. Box69, Yaounde,
Cameroon
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
... In the tropical forests, we monitored the herbs and smaller shrubs using security cameras (Vivotek IB8367) with a target to record ten individuals of each plant species and each plant individual for two days. Trees and shrubs, which did not fit the camera field view, were observed personally with the target to observe eight individuals of each plant species each for eight hours (for more details, see Janeček et al. 2022). In montane grasslands, where the vegetation is much more open, we observed more plants at one observation point. ...
... This was true mainly in the dry season, when the observed niche overlap was notably high, although not significantly greater than that predicted by the null model. In the MF, during the dry season, trees Astropanax abyssinicum, Nuxia congesta, and Syzigium staudtii were the plants most frequently visited by C. preussi ( Fig. 1; Janeček et al. 2022). In the SF, during the dry season, the most visited plants were trees Syzigium sp. and Tabernaemontana ventricosa. ...
... Nevertheless, precise trait matching in the entire sunbird-plant network seems to be obscured by asymmetry in the relationship between ornithophilous plants and sunbirds ). Sunbirds do not care much and visit many other non-specialized plants in both the wet and dry seasons Janeček et al. 2022). Moreover, the absence Results of chi-square tests comparing the observed number of visits with the predicted number of visits when the female/male visitation ratio will be 1. ...
Article
Full-text available
Competition for nectar is expected to result in feeding niche differentiation. Here, we targeted the sexually size-dimorphic Volcano Sunbird ( Cinnyris preussi ) on Mount Cameroon. We investigated whether males and females feed on different plant species, whether females with shorter bills than males prefer shorter flowers, and whether larger dominant males visit more energetic flowers that produce higher amounts of nectar sugars. We also asked whether feeding niches were spatially separated along the elevation gradient and whether this separation varied between the two contrasting seasons. We collected data on the frequency of visits to individual plant species and analyzed the male-to-female ratios in the mist-netted dataset. In addition, we estimated production of nectar sugar in individual habitats and seasons. Despite the large dataset collected, encompassing 6476 bird–plant interactions, our findings did not provide evidence of differences in the spectra of the visited plant species. In addition, females did not visit flowers with shorter tubes, nor did males visit flowers that produced higher amounts of sugars. However, we observed a sex-specific dispersion of sunbirds during the wet season. During the dry breeding season, both males and females feed mainly in nectar-rich montane and submontane forests. In the wet season, the production of nectar sugar in these habitats decreased dramatically, and females largely disappeared. In contrast, female activity increased in the lowest and highest parts of the altitudinal range. Our findings on elevational movements are important in the current context, in which species face potential threats from habitat destruction and climate change.
... Therefore, Cyanomitra olivacea selectively foraged on the plant species with the largest amount of nectar. Similarly, Janeček et al. (2022) found that sunbirds were more specialized at lower elevations where Cyanomitra olivacea is found. ...
... on sunbird-plant interactions were collected by observation of individual plant specimens (used inJaneček et al., 2022). ...
Article
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The drivers behind plant-pollinator interactions still need to be fully understood. Previous research has suggested that observed interactions result from either neutral interaction between species based on their abundance or from niche-based processes, which are reflected in the adaptations of both plants and pollinators. Furthermore, the importance of both scenarios can differ depending on seasonal dynamics translated into differences in resource availability. Extensive research has been conducted on New-World hummingbirds (Trochilidae), whereas much less is known about Old World nectar-feeding sunbirds (Nectariniidae). Our study aimed to explore whether sunbird-plant interactions are explained by neutral and/or niche-based processes. Thus, we tested the effects of abundance, morphology, and nectar sugar content on the observed interactions and the link between sunbirds and plant traits. Moreover, we explored the effects of robbing on these mechanisms. Finally, we investigated the partitioning of bird pollination niches, based on floral traits. We used a dataset of sunbird-plant interactions collected at Mount Cameroon during two seasons (dry and wet seasons). Our study shows that sunbird-plant interactions are influenced by both neutral and niche-based processes. Neutral processes and nectar reward were the main mechanisms underlaying the observed interactions in the dry season. However, as more ornithophilous plants bloom during the rainy season, morphological trait-matching becomes more important. We found a correlation between bill length and floral tube dimensions and observed niche overlap among the sunbird species. Considering this and other research, we suggest that plant-pollinator interactions are influenced by a combination of both neutral and niche-based processes.
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Aim Temporal dynamics of biodiversity along tropical elevational gradients are unknown. We studied seasonal changes of Lepidoptera biodiversity along the only complete forest elevational gradient in the Afrotropics. We focused on shifts of species richness patterns, seasonal turnover of communities and seasonal shifts of species’ elevational ranges, the latter often serving as an indicator of the global change effects on mountain ecosystems. Location Mount Cameroon, Cameroon. Taxon Butterflies and moths (Lepidoptera). Methods We quantitatively sampled nine groups of Lepidoptera by bait‐trapping (16,800 trap‐days) and light‐catching (126 nights) at seven elevations evenly distributed along the elevational gradient from sea level (30 m a.s.l.) to timberline (2,200 m a.s.l.). Sampling was repeated in three seasons. Results Altogether, 42,936 specimens of 1,099 species were recorded. A mid‐elevation peak of species richness was detected for all groups but Eupterotidae. This peak shifted seasonally for five groups, most of them ascending during the dry season. Seasonal shifts of species’ elevational ranges were mostly responsible for these diversity pattern shifts along elevation: we found general upward shifts in fruit‐feeding butterflies, fruit‐feeding moths and Lymantriinae from beginning to end of the dry season. Contrarily, Arctiinae shifted upwards during the wet season. The average seasonal shifts of elevational ranges often exceeded 100 m and were even several times higher for numerous species. Main conclusions We report seasonal uphill and downhill shifts of several lepidopteran groups. The reported shifts can be driven by both delay in weather seasonality and shifts in resource availability, causing phenological delay of adult hatching and/or adult migrations. Such shifts may lead to misinterpretations of diversity patterns along elevation if seasonality is ignored. More importantly, considering the surprising extent of seasonal elevational shifts of species, we encourage taking account of such natural temporal dynamics while investigating the global climate change impact on communities of Lepidoptera in tropical mountains.
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Sunbirds play a major role in the pollination of Old World nectivorous plants. However, with the exception of the Cape Floristic Region there is a major knowledge gap around African nectivore interaction networks—a stark contrast from the abundance of neotropical hummingbird–plant networks. Here, we describe a sunbird pollen transfer network (PTN) which we use in conjunction with a sunbird flower visitation network (FVN) to explore levels of sunbird specialization within an Afromontane forest habitat. Both networks were generalized compared with similar‐sized hummingbird networks, reflecting the wide range of flower types visited, the generalist diet, and bill characteristics of sunbirds. Three sunbird species from the genus Cinnyris accounted for 85% of flower visits and 77% of all pollen transported. Of the 17 plant species across both networks, 15 are predominantly pollinated by insects while Anthonotha noldeae (Fabaceae–Caesalpinioideae) and Globimetula braunii (Loranthaceae) depend on sunbirds for seed set. Sunbird species average bill lengths varied between 14.5 mm (the variable sunbird) and 23.6 mm (the Green‐headed Sunbird), but, while more pollen was carried on longer bills, we found no evidence for a relationship between bill length and type of flower visited. Both networks were nested. Some specialization was observed in both networks although this does not appear to be driven much by sunbird–flower trait matching. Overall, our results suggest that in contrast to nectivores elsewhere, factors such as phenology and/or environment, rather than morphology, may play important roles in limiting potential sunbird–flower interactions and need further investigation.
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The effect of perching versus hovering on the drinking speed of the Cameroon Sunbird Cyanomitra oritis and the Northern Double-collared Sunbird Cinnyris reichenowi was investigated in a cage experiment. Our results support the hypothesis that the shorter handling times on flowers during hovering compared to perching are largely caused by an increase in drinking speed.
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Aim To test if tree species richness and forest structure drive spatial variation in avian communities along a tropical elevation gradient and to present information about the role of detailed forest parameters. Location A 2,000‐m long elevational gradient of tropical forest on Mt. Cameroon, west‐central Africa. Taxon Birds and trees. Methods We performed bird censuses and vegetation mapping at the same plots across six forested sites at elevations of 350, 650, 1,100, 1,500, 1,850, and 2,200 m a.s.l., with 16 plots per elevation. We tested the effects of elevation, forest structure and tree diversity on the species richness, functional diversity and β‐diversity of birds (Bray–Curtis dissimilarity). We used conditional inference trees based on random forests (RF) to investigate these relationships across all elevation sites as well as within elevations. Results Both tree and bird species richness declined monotonically with elevation. Vegetation structure correlated with elevation, and all vegetation attributes significantly differed among elevations. The RF explained 70% of the variance in avian species richness, with the most important predictors being elevation, proportion of dead trees, tree species richness and herb layer coverage. We found that elevation (and shrub layer) was a particularly important predictor of avian functional diversity. We identified no important predictor of bird species richness after standardization within elevations, and the proportion of dead trees was the sole important predictor of functional diversity. Within‐elevation β‐diversity in avian community composition was determined by the dissimilarity of the tree community and differences in leaf area index, solar radiation and spatial distance. The functional dissimilarity was best explained by leaf area index. Main conclusions Apart from elevation itself, spatial distance even within elevations correlated with compositional and functional variation among avian assemblages. Forest structural traits can have a significant influence on distribution of birds. Thus, gaps in the spatial distribution of species such as along elevations might be caused by fine‐scale recognition of suitable habitats.