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An Australian painted lady (Vanessa kershawi) feeding on nectar through its long proboscis

In zoology, a nectarivore is an animal which derives its energy and nutrient requirements from a diet consisting mainly or exclusively of the sugar-rich nectar produced by flowering plants.

Nectar as a food source presents a number of benefits as well as challenges. It is essentially a solution of (as much as 80%) the simple sugars sucrose, glucose and fructose, which are easily ingested and digested, representing a rich and efficient source of nutrition. This solution is often diluted either by the plant that produces it or by rain falling on a flower and many nectarivores possess adaptations to effectively rid themselves of any excess water ingested this way.

However, nectar is an incomplete source of nutrition. While it does contain proteins and amino acids,[1] these are found in low quantities, and it is severely deficient in minerals and vitamins.[2] Very few organisms consume nectar exclusively over their whole life cycle, either supplementing it with other sources, particularly insects (thus overlapping with insectivores) or only consuming it exclusively for a set period.[3] Many species are nectar robbers or nectar thieves, performing no pollination while still consuming nectar. Many species are both nectar robbers and pollinators, depending on the plant species they encounter.

Nectar is produced by flowering plants to attract pollinators to visit the flowers and transport pollen between them. Flowers often have specialized structures that make the nectar accessible only for animals possessing appropriate morphological structures, and there are numerous examples of coevolution between nectarivores and the flowers they pollinate. For example, hummingbirds and hawkmoths have long narrow beaks that can reach nectar at the bottom of long tubular flowers.[4][5]

The majority of nectar feeders are insects or birds, but instances can also be found in other animal groups.

Insects

[edit]
An Eastern carpenter bee (Xylocopa virginica) pierces the corolla to feed from a daffodil (Narcissus sp.)

Nectarivory is extremely common in insects. Key families with large proportions of nectarivores include the Coleoptera, Lepidoptera, Diptera, Hymenoptera and Hemiptera. Some, but not all, are also pollinators: others engage in nectar robbing by avoiding the reproductive organs of plants altogether, particularly those with deep corollas, by piercing into the base of the flower to reach the nectary directly, such as carpenter bees and secondarily honey bees (who consume nectar from holes made by others),[6] as well as ants, who frequently consume nectar and pollen where available despite actively inhibiting germination of pollen at the flowers they visit to the detriment of the plant.[7]

Two Spot swordtail butterflies (Graphium nomius) mud puddling for minerals

Nectar-feeding insects gain enough water from nectar to rarely need to drink, though adult butterflies and moths may engage in puddling in order to obtain dissolved substances not abundant in nectar, particularly salts and amino acids.[8] Some flying nectarivores, particularly larger bees, do not lose enough water by evaporation while on the wing to offset their high intake due to nectar-feeding, as well as water produced metabolically while flying. They must excrete while on the wing to prevent water loading, and may wait at the nest entrance to evaporate off some of their water load before flying out.[9]

Arachnids

[edit]

There is evidence that some spiders, though normally thought to be exclusively carnivorous, consume nectar indirectly by consuming nectarivorous insects, and/or directly from flowers. This behavior is thought to be more common among spiders that live among foliage. A few make nectar their primary food source, such as Bagheera kiplingi, a member of the jumping spiders,[10][11][12] while others such as the crab spiders, feed more rarely and opportunistically. None of the spider groups observed feeding on nectar build webs, they are all wandering species.[13]

Birds

[edit]
A female ruby-throated hummingbird (Archilochus colubris) feeds on nectar from a sunflower (Helianthus annuus)

Nectar-feeding is widespread among birds, but no species consumes nectar exclusively. Most combine it with insectivory for a mixed diet. Of particular interest are three lineages of specialized nectarivorous birds: the hummingbirds (Trochilidae), sunbirds (Nectariniidae) and honeyeaters (Meliphagidae). These groups have adapted to permit a nectar-central diet, showing higher activity of digestive enzymes which break down sugars, higher rates of absorption of sugars, and altered kidney function. To maintain flight a bird must rapidly excrete much of the water content of the nectar it consumes. A hummingbird's kidneys are capable of rapidly producing large quantities of hyposmotic urine i.e. urine containing a lower concentration of dissolved substances than the blood.[14] Some other bird groups have one or more similar specializations – for instance, the Lories, one group of Australasian parrots within the larger parrot family Psittacidae, possess similar digestive modifications.[15] These are examples of parallel evolution. The Hawaiian honeycreepers have several species adapted to feed on nectar. The Hawaiian tree Metrosideros polymorpha is heavily dependant on the pollination of the more or less nectarivorous honeycreepers.[16]

Mammals

[edit]
A grey-headed flying fox (Pteropus poliocephalus) feeds on nectar, its face covered with yellow pollen

Many species of bat feed on nectar, their lifestyle similar to that of nectarivorous birds. In the Americas there is significant overlap between flowers pollinated by bats and hummingbirds – both need similarly-composed nectar to keep up energy-intensive hovering flight. In this part of the world there is particularly close association between some species of columnar cacti and bat species, who provide pollination in exchange for nectar with composition matching their nutritional needs.[17] Nectarivorous bats might be at particular risk of extinction due to their reliance on particular species of flowering plants.[18]

A single marsupial species, the honey possum, feeds on nectar and pollen exclusively. It raises fewer young which grow more slowly than other marsupials of its size, because of the time-consuming effort of nectar-drinking from many flowers to support itself. It may spend periods in deep sleep to reduce its need for food, and shows the typical nectarivore adaptations for excess water-removal.[19]

See also

[edit]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Nectarivore

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A nectarivore is an animal that derives its primary energy and nutrients from a diet consisting mainly or exclusively of , the sugar-rich liquid secreted by the flowers of angiosperms to attract pollinators. This feeding strategy, known as nectarivory, has evolved independently multiple times across diverse animal lineages, enabling these organisms to exploit floral resources while often serving as key pollinators in ecosystems. Nectarivores exhibit remarkable physiological and morphological adaptations suited to their liquid diet, including elongated tongues or proboscides for accessing deep floral nectaries, rapid digestive systems to process high-sugar intake, and specialized sensory capabilities such as ultraviolet vision in insects or hovering flight in birds and bats. Among the most prominent groups are insects, which form the largest cohort of nectarivores—including bees (Apidae), butterflies (Lepidoptera), and hawkmoths (Sphingidae)—and play a foundational role in plant-pollinator mutualisms by transferring pollen during feeding. Vertebrate nectarivores include around 800 bird species, such as hummingbirds (Trochilidae), sunbirds (Nectariniidae), and honeyeaters (Meliphagidae), which dominate in tropical and subtropical regions and promote high levels of plant genetic diversity through their pollination services. Nectarivorous bats, comprising approximately 70 species primarily in the families Phyllostomidae (Neotropics) and Pteropodidae (Old World), feature specialized tongue structures—like brush-tipped papillae or grooved pumping mechanisms—for efficient nectar extraction, and they pollinate hundreds of plant species, particularly nocturnal flowers. Less common are mammalian nectarivores, such as certain marsupials (e.g., honey possums Tarsipes rostratus) and lemurs, which supplement nectar with pollen or insects to meet protein needs. Ecologically, nectarivores drive evolution, , and community structure by exerting selective pressure on floral traits like color, shape, and composition, while their behaviors enhance production, rates, and resilience. For instance, bird-pollinated often display red, tubular flowers with dilute , adapted to the birds' high-energy demands and mobility, resulting in nearly twice the number of mating partners per compared to insect-pollinated . However, nectarivory also presents challenges, such as nutritional imbalances from low protein content in , leading many nectarivores to consume , , or other supplements. Overall, these interactions underscore the intricate balance between nectarivores and , supporting in diverse habitats worldwide.

Definition and Characteristics

Definition

A nectarivore is an animal that derives a significant portion, often the , of its energy and nutritional requirements from , a sugar-rich secreted by floral nectaries as a reward for pollinators. This dietary specialization distinguishes nectarivory from more generalist feeding strategies, emphasizing reliance on nectar's high-carbohydrate content for activities such as flight and . Nectar typically comprises 50–80% by weight, with sugars accounting for 10–70% (w/w) and forming 80–90% of the dry matter, primarily as , , and in varying ratios depending on plant species and environmental factors. It also includes trace amounts of (0.19–12.7 mM total, encompassing all 20 proteinogenic types, often dominated by non-essential ones like and ), as well as secondary compounds such as phenolics, alkaloids, and . While these sugars serve as an efficient source, nectar generally lacks substantial proteins and , which occur only in low concentrations and are not primary nutritional contributors. Unlike pollinivores, which primarily consume pollen for protein, nectarivores focus specifically on nectar intake, without necessarily exploiting or other floral structures. The concept of nectarivory emerged in 19th-century and , with foundational observations by on the mutualistic systems where rewards attract and sustain animal visitors. The term "nectarivore" itself was coined later, with its earliest recorded use in 1967 within ecological literature.

Adaptations

Nectarivores exhibit a suite of morphological adaptations that facilitate access to nectar hidden within floral structures. Insects often possess elongated proboscises, which function as siphons to draw nectar from deep corollas, as seen in long-proboscid flies where the proboscis length corresponds to specific flower tube dimensions for efficient feeding. In birds and mammals, tubular bills and extensible tongues enable precise probing; for instance, the hummingbird tongue features a reversible, forked tip that traps nectar through elastic expansion and contraction during lapping, allowing uptake without reliance on capillary action. Physiological adaptations address the challenges of nectar's high and sugar composition. Specialized kidneys in birds like hummingbirds permit rapid excretion of dilute , processing 3–5 times their body weight in daily to manage osmotic loads from intake. Gut microbiomes in species such as sunbirds contribute to detoxifying secondary metabolites in , enhancing nutrient absorption. Behavioral adaptations optimize energy acquisition while minimizing risks. Hovering flight in birds allows stationary access to pendant flowers, reducing the need to and enabling rapid visits to multiple blooms. Nocturnal feeding in bats synchronizes with flower peaks at night, exploiting less competitive resources under cover of darkness. Certain nectarivores employ nectar-robbing tactics, such as piercing corolla bases to extract rewards without contacting reproductive parts, thereby avoiding duties while securing food. Nectar’s low protein content poses nutritional challenges, prompting supplementation strategies like pollen consumption in birds to meet amino acid needs during breeding seasons. This diet also demands high metabolic rates for energy processing; hummingbirds, for example, sustain heart rates up to 1,200 beats per minute to fuel rapid digestion and activity. Convergent evolution underscores these adaptations across taxa, with unrelated groups developing analogous structures for similar ecological niches. Sunbirds and hummingbirds, despite phylogenetic distance, share elongated bills and brush-tipped tongues for nectar extraction, reflecting parallel responses to selective pressures from tubular flowers.

Ecological and Evolutionary Aspects

Ecological Role

Nectarivores play a central role in mutualism, where offer nectar as a reward to attract these animals, which in turn facilitate transfer between flowers. This interaction is essential for the of many flowering , with animal pollinators, including nectarivores, responsible for approximately 90% of angiosperm globally. In some habitats, such as tropical forests, this dependency exceeds 94%, underscoring the disproportionate influence of nectarivores on in hotspots. Through their activities, nectarivores provide key services that bolster plant diversity and support . They maintain floral communities by enabling and preventing , which enhances overall and resilience in ecosystems. In , nectarivores contribute to the of crops like almonds, where managed and wild pollinators ensure fruit set and yield stability, accounting for a significant portion of global food production volume—up to 35% for animal-pollinated crops. Additionally, nectarivorous birds serve as indicators of , as their populations are highly sensitive to alterations, signaling broader declines. Nectarivores engage in various interactions that shape community dynamics, including for limited resources among , which can influence behaviors and resource partitioning. Predation risks also arise at floral sites, such as spiders ambushing nectar-feeding or birds at flowers, potentially disrupting efficiency. Indirectly, nectarivores contribute to by promoting fruit production through successful , as pollinated plants develop fruits that attract frugivores for transport. Habitat fragmentation and pesticide use pose major threats to nectarivores, reducing available sources and directly harming populations through exposure and loss of areas. In tropical ecosystems, where nectarivores drive over 90% of , these pressures exacerbate declines, threatening both floral diversity and associated food webs. Conservation efforts emphasize protecting connected habitats to sustain nectarivore services. Recent post-2020 research highlights eco-evolutionary feedbacks, where herbivores indirectly boost production by altering dynamics and fitness trade-offs.

Evolutionary History

Nectarivory first emerged during the Middle-Late , approximately 240–201 million years ago, associated with 'flowers,' with significant diversification during the period (~145–66 million years ago) coinciding with the rapid radiation of angiosperms that provided new floral resources for animal exploitation. This dietary specialization initially developed among , with early evidence from pollen-feeding beetles transitioning to nectar consumption as gymnosperm-associated lineages shifted to angiosperm hosts around the mid-. Fossil records indicate that insect nectarivores, such as long-proboscid flies, predated vertebrate adopters by millions of years, leveraging specialized mouthparts for accessing in primitive flowers. The of nectarivory has been tightly linked to coevolutionary dynamics between animals and , where flower morphology adapted to match traits, such as the development of long-tubed corollas suited to specialized feeders with elongated proboscises or bills. Concurrently, chemistry evolved to selectively attract or deter specific taxa, incorporating sugars, , and secondary metabolites that align with the digestive capabilities of target pollinators while discouraging non-pollinators. For instance, sucrose-dominant nectars in many hummingbird-pollinated flowers reflect reciprocal adaptations in avian sucrase enzyme activity, exemplifying convergent across distant lineages. Convergent evolution has driven independent origins of nectarivory in multiple groups, including birds like New World hummingbirds (Trochilidae) and sunbirds (Nectariniidae), which separately developed hovering flight and tubular tongues despite phylogenetic distance. Similarly, among mammals, nectar-feeding bats (e.g., glossophagine species) and marsupials such as the (Tarsipes rostratus) exhibit parallel traits like elongated, brush-tipped tongues and enhanced nectar-processing enzymes, arising from unrelated ancestral diets. In parrots, parallel adaptations to nectarivory, including bill shape and gut morphology, show strong phenotype-environment correlations that facilitated diversification in lories and lorikeets. Diversification of nectarivory in involved multiple independent transitions, such as from carnivorous ancestors in beetles, where mid-Cretaceous fossils reveal mouthparts specialized for both and intake, marking a shift toward angiosperm reliance. Fossil evidence from further documents ancient nectar-feeding with siphonate proboscides dating back to the , underscoring repeated innovations in insect lineages during the angiosperm terrestrial revolution. Recent research from 2024 highlights of nectaries in ferns, which independently recruited bodyguards via sugary rewards during the , paralleling angiosperm strategies and expanding -animal mutualisms beyond flowers. Eco-evolutionary models further demonstrate how herbivores indirectly shape evolution by suppressing growth, thereby selecting for increased nectar production to bolster attraction and defense. These insights, drawn from phylogenetic and dynamic simulations, reveal ongoing feedbacks that refine nectar composition in response to multi-trophic pressures.

Nectarivores by Taxonomic Group

Insects

Insects constitute the most diverse group of nectarivores among animals, encompassing the vast majority of known pollinators and nectar-feeding , with estimates exceeding 200,000 primarily within this class. These insects play a critical ecological role, contributing to the of approximately 35% of global food crops through their foraging behaviors. The primary orders engaging in nectarivory include , , Diptera, and Coleoptera, each exhibiting specialized mouthpart adaptations that facilitate nectar extraction from flowers. In the order Hymenoptera, bees are prominent nectarivores, often equipped with pollen baskets (corbiculae) on their hind legs for simultaneous collection of nectar and pollen, enhancing their efficiency as pollinators. Lepidopterans, such as butterflies and moths, possess a coiled proboscis that uncoils to access nectar, with lengths reaching up to 28 cm in some species to exploit deep floral tubes. Dipterans, or flies, typically feature short, lapping mouthparts suited for consuming nectar from shallow or exposed flowers, allowing them to act as generalist feeders across diverse plant communities. Coleopterans, including beetles, rely on chewing mandibles modified with cutting edges and teeth to access nectar, often feeding on both floral rewards and plant tissues in a less specialized manner. Notable examples illustrate the behavioral diversity of insect nectarivory. The eastern carpenter bee (Xylocopa virginica) exemplifies generalist feeding, targeting a wide range of flowers for nectar while employing powerful mandibles to pierce corollas. In Lepidoptera, the monarch butterfly (Danaus plexippus) depends heavily on nectar to fuel its long-distance migration, converting sugars into lipid reserves essential for the journey spanning thousands of kilometers. Hawkmoths of the family Sphingidae demonstrate nocturnal pollination strategies, hovering to feed on nectar from pale, fragrant flowers using an elongated proboscis, thereby facilitating reproduction in night-blooming plants. Unique aspects of insect nectarivory include nectar-robbing behaviors, where individuals like pierce the base of flowers to extract without contacting reproductive structures, thus bypassing . Social species within , such as , process collected through regurgitation and enzymatic activity to produce , a concentrated energy store for colony sustenance. Nectarivory has evolved independently across insect lineages, often representing a dietary shift from ancestral habits like predation or foliage consumption to exploit floral resources.

Arachnids

Nectarivory among arachnids is predominantly documented in spiders (order Araneae), where it supplements a primarily carnivorous diet by providing carbohydrates, amino acids, and hydration. This behavior has been observed in over 60 spider species across 10 families, with floral and extrafloral nectar being the most common plant-derived food source. Unlike the specialized nectarivory seen in many insects, spider nectar feeding is typically opportunistic, aiding survival during prey shortages or in resource-poor habitats such as arid regions. A notable example is the Bagheera kiplingi (family Salticidae), which exhibits an extreme shift toward herbivory; in Mexican populations, plant materials including extrafloral from Vachellia trees and Beltian bodies constitute over 90% of its diet. Other salticids, such as Phidippus and Menemerus , regularly visit flowers to imbibe , with laboratory tests confirming this capability in 90 and field observations in at least 31. Crab spiders (family ), including genera like Misumena and Thomisus, position themselves on flowers to ambush pollinators but also directly consume and during periods of low prey availability, enhancing their endurance on inflorescences. Spiders access nectar primarily through direct licking with their chelicerae, often pressing mouthparts against nectaries or using palps and forelegs to extract droplets, as observed in salticids visiting up to 80 flowers per hour guided by olfactory cues. Indirect nectar intake occurs via predation on nectar-laden insects like bees or flies, though this is incidental to their carnivorous habits. In arid environments, nectar's high water content (often 50-80%) provides essential hydration, allowing species like wandering hunters in the family Anyphaenidae to persist in dry habitats. This dietary flexibility underscores broader opportunism in spiders, enabling physiological adaptations to variable food resources without a full transition from carnivory.

Birds

Avian nectarivores represent a diverse group of birds specialized for feeding on floral , primarily within several and non-passerine families. The most prominent is the Trochilidae, or hummingbirds, comprising over 360 species exclusively in the , from to , where they dominate nectar-feeding niches through hovering flight and long bills adapted for tubular flowers. In the , the Nectariniidae (sunbirds) are key, with around 140 species distributed across , southern Asia, and parts of , featuring slender, curved bills for accessing deep corollas. The Meliphagidae (honeyeaters), with approximately 180 species, prevail in , including , , and Pacific islands, often perching to probe blossoms with brush-tipped tongues. Additionally, certain members of the (parrots), particularly the Loriinae subfamily of lorikeets, have independently evolved nectarivory, with about 50 species in and nearby regions exhibiting specialized papillae-covered tongues for lapping and pollen. Representative examples illustrate these adaptations in action. The (Archilochus colubris), common in eastern , visits up to 2,000 flowers daily to meet its energy demands, pollinating plants like bee balm and cardinal flower in the process. In , the (Phylidonyris novaehollandiae) forages aggressively on native flowers such as , using its tubular tongue to extract while also consuming insects for protein. The rainbow lorikeet (Trichoglossus moluccanus), widespread in eastern and introduced elsewhere, employs its unique brush-like tongue with projecting papillae to collect and from eucalypts and other trees, often in noisy flocks. These birds exhibit unique behavioral and physiological traits suited to nectar dependence. Many defend nectar-rich patches territorially, with hummingbirds engaging in aerial chases to monopolize feeders or flower clusters, ensuring exclusive access to high-energy resources. Their metabolic rates are extraordinarily high; hummingbirds, for instance, must consume nectar equivalent to half their body weight daily to offset overnight losses of up to 10% of mass during , a state that reduces energy expenditure. Parallel evolution is evident between New World hummingbirds and Old World sunbirds or honeyeaters, with convergent traits like extensible tongues and specialized bills arising independently to exploit similar floral resources.

Mammals

Mammalian nectarivores are primarily found among bats in the order Chiroptera and certain marsupials in the infraclass Marsupialia, with bats representing the most diverse and specialized group. In Chiroptera, nectar-feeding species are concentrated in the subfamily Glossophaginae within the family Phyllostomidae, predominantly in the Neotropics, where they exhibit morphological and behavioral adaptations for nocturnal foraging. Among marsupials, the (Tarsipes rostratus) is an obligate nectarivore endemic to southwestern , while some pygmy possums in the Cercartetus opportunistically consume and . These mammals play crucial ecological roles as pollinators, supplementing their high-energy diets with and to meet nutritional needs. Nectarivorous bats, such as those in Glossophaginae, rely on echolocation to locate flowers in the dark, emitting high-frequency calls to detect sources from distances up to several meters. They possess elongated snouts and s fringed with hair-like papillae that function like mops to extract efficiently from deep corollas, enhancing feeding rates in sympatric species. A prominent example is the lesser long-nosed (Leptonycteris yerbabuenae), a migratory species that travels over 1,000 kilometers annually from to the southwestern United States, pollinating columnar cacti and agaves along a seasonal corridor. In contrast, the (Pteropus poliocephalus), a pteropodid from eastern , combines and from eucalypts with in its diet, using its brush-tipped to lap up floral rewards while roosting in large colonies. Among marsupials, the (Tarsipes rostratus) is uniquely adapted as the only non-flying with an exclusive diet of and , featuring a long, extensible tongue with a brush tip for collecting rewards from diverse proteaceous and myrtaceous flowers, and specialized kidneys to handle high water intake. Pygmy possums, such as Cercartetus species, are opportunistic feeders that prioritize from eucalypts and banksias when available, supplementing with for protein, and use to conserve energy during nectar shortages. These adaptations allow them to thrive in heathlands and woodlands, though their small size (10-40 grams) imposes high metabolic demands. Certain , particularly lemurs in , also engage in nectarivory. Species like the (Varecia variegata) feed on from large flowers such as the traveler's palm (Ravenala madagascariensis), using long tongues to access rewards without damaging blooms, thereby serving as important pollinators. This behavior supplements their primarily frugivorous diet and highlights in mammalian nectar feeding. Conservation challenges for mammalian nectarivores are acute, driven by habitat loss and degradation of nectar-producing plants. Over 20 of nectar-feeding bats are classified as endangered or vulnerable globally, primarily due to and agricultural expansion reducing floral resources in the Neotropics and . These bats pollinate more than 300 plant , including economically vital crops like (used for production) and , underscoring their irreplaceable . The faces in southwestern , though not yet listed as endangered, its populations are declining from urban development and altered fire regimes. Efforts to mitigate include restoration and protected corridors to sustain these vital pollinators.

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