Nectarivore

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.

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

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 services to a plant while still consuming nectar.

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. Bats, meanwhile, visit open flowers where the nectar is not as deeply hidden.[4][5]

Nectarivorous species

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

Birds

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.[6] 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.[7] These are examples of parallel evolution. The Hawaiian honeycreepers have several species adapted to feed on nectar. Especially important is the Hawaiian tree Metrosideros polymorpha which is heavily depending on the pollination of the more or less nectarivourus honeycreepers.[8]

Insects

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),[9] 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.[10]

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.[11] 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.[12]

Arachnids

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,[13][14][15] 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.[16]

Mammals

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. There is significant overlap between flowers pollinated by bats and hummingbirds – both need similarly-composed nectar to keep up energy-intensive hovering flight. In the Americas there is particularly close association between some species of columnar cacti and bat species, who provide pollination services in exchange for nectar with composition matching their nutritional needs.[17] Nectarivorous bats are 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, which limits its available energy. It may spend periods in deep sleep to reduce wasted energy and shows the typical nectarivore adaptations for excess water-removal.[19]

• detritivore
• Palynivore
• Frugivore
• Herbivore

References

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2. Gartrell B (2000). "The Nutritional, Morphologic, and Physiologic Bases of Nectarivory in Australian Birds". Journal of Avian Medicine and Surgery. 14 (2): 85–94. doi:10.1647/1082-6742(2000)014[0085:TNMAPB]2.0.CO;2.
3. Nicolson S, Fleming P (2014). "Drinking problems on a 'simple' diet: physiological convergence in nectar-feeding birds". J Exp Biol. 217 (7): 1015–23. doi:10.1242/jeb.054387. PMID 24671960.
4. Wiens, John A. (1992). The Ecology of Bird Communities: Processes and variations. p. 336. ISBN 9780521426350.
5. Richard, A.J. (1997). Plant breeding systems (Second ed.). Chapman & Hall. ISBN 978-0-412-57440-5.
6. Nicolson S, Fleming P (2014). "Drinking problems on a 'simple' diet: physiological convergence in nectar-feeding birds" (PDF). J Exp Biol. 217 (7): 1015–23. doi:10.1242/jeb.054387. PMID 24671960.
7. Schweizer M, Güntert M, Seehausen O, Leuenberger C, Hertwig S (2014). "Parallel adaptations to nectarivory in parrots, key innovations and the diversification of the Loriinae". Ecol Evol. 4 (14): 2045–7758. doi:10.1002/ece3.1131. PMC 4130445. PMID 25165525.
8. Pratt, H. Douglas; Conant, Shelia (2005-05-12). The Hawaiian Honeycreepers: Drepanidinae. OUP Oxford. ISBN 978-0-19-854653-5.
9. Dedej S, Delaplane K (2005). "Net energetic advantage drives honey bees (Apis mellifera) to nectar larceny in Vaccinium ashei Reade". Behavioral Ecology and Sociobiology. 57 (4): 398–403. doi:10.1007/s00265-004-0852-z.
10. Hull D, Beattie A (1988). "Adverse effects on pollen exposed to Atta texana and other North American ants: implications for ant pollination". Oecologia. 75 (1): 153–155. doi:10.1007/BF00378829. PMID 28311849.
11. Beck J, Mahlenberg E, Fiedler K (1999). "Mud-puddling behavior in tropical butterflies: in search of proteins or minerals?". Oecologia. 119 (1): 140–148. doi:10.1007/s004420050770. PMID 28308154.
12. Nicolson S (2009). "Water homeostasis in bees, with the emphasis on sociality". J Exp Biol. 212 (3): 429–434. doi:10.1242/jeb.022343. PMID 19151218.
13. Mihai, Andrei (Oct 13, 2009). "First (mainly) vegetarian spider found". ZME Science.
14. Jackson R, Pollard S, Nelson X, Edwards G, Barrion A (2001). "Jumping spiders (Araneae: Salticidae) that feed on nectar". J. Zool. 255 (1): 1469–7998. doi:10.1017/S095283690100108X.
15. Meehan J, Olson E, Reudink M, Kyser T, Curry R (2009). "Herbivory in a spider through exploitation of an ant–plant mutualism". Curr. Biol. 19 (19): 892–893. doi:10.1016/j.cub.2009.08.049. PMID 19825348.
16. Taylor R, Pfannenstiel R (2008). "Nectar Feeding by Wandering Spiders on Cotton Plants". Environ. Entomol. 37 (4): 996–1002. doi:10.1093/ee/37.4.996. PMID 18801266.
17. Nassar J, Ramirez N, Linares O (1997). "Comparative pollination biology of Venezuelan columnar cacti and the role of nectar-feeding bats in their sexual reproduction". Am. J. Bot. 84 (7): 918–927. doi:10.2307/2446282. JSTOR 2446282. PMID 21708646.
18. Arita H, Santos-Del-Prado K (1999). "Conservation Biology of Nectar-Feeding Bats in Mexico". J. Mammal. 80 (1): 31–41. doi:10.2307/1383205. JSTOR 1383205.
19. Richardson K, Wooller R, Collins B (1986). "Adaptations to a diet of nectar and pollen in the marsupial Tarsipes rostratus (Marsupialia: Tarsipedidae)". J. Zool. 208 (2): 285–297. doi:10.1111/j.1469-7998.1986.tb01515.x.