Enchenopa binotata complex

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Enchenopa binotata complex
Two-marked Treehopper (Enchenopa binotata) - Guelph, Ontario.jpg
Scientific classification
Kingdom:
Animalia
Phylum:
Arthropoda
Class:
Insecta
Order:
Hemiptera
Family:
Membracidae
Genus:
Enchenopa
Species complex:
E. binotata
Binomial name
Enchenopa binotata
Map E.bin.jpg
Map showing the distribution of E. binotata [1]

Enchenopa binotata (Hemiptera: Membraciade) is a complex of multiple species found mostly in Eastern North America, [2] [3] [4] but have also been reported in Central America. [5] They are commonly referred to as treehoppers and are sap-feeding insects. [2] The species in the complex look similar to each other in morphology, [6] but are identified as different species by the host plant they occupy.

Contents

Biology

Morphology

Nymphs (juveniles) of E. binotata start out to be ≤1mm with gray and black coloration. Nymphs have 5 instars until they molt into adulthood, which can take 3–4 weeks. [6] As adults, they can range from 7-9mm in size. [7] and have two yellow markings on their back. Their species name is derived from these two markings; bi-, meaning "two", -notata, meaning "to mark". They form thornlike structures on their head called a pronotum. These treehoppers are true bugs, belonging to the order Hemiptera, which all share a common mouth morphology; sucking mouthparts. [8]

Vibrational communication

Male E. binotata 'Ptelea' treehopper from an Illinois population signaling at 24 °C
E. binotata male signal that contains 2 signals in 1 bout with 2 pulses each Enchenopa binotata Ptelea signal.jpg
E. binotata male signal that contains 2 signals in 1 bout with 2 pulses each

Male E. binotata treehoppers make substrate-borne vibrations on the stems, petioles, and leaves of their host plants that travel throughout the plant. Females detect these vibrational signals with specialized structures on their legs and they also respond through the plant. [10] Male signals are more complex than female responses. Male and female signals are tonal, but females respond with grunt-like sounds that are at a lower frequency than the males. Females have specific species preferences, and prefer signals that are close to these conspecific frequencies, but frequencies can change with temperature fluctuations. [11] [12] Despite these fluctuations, females are able to distinguish their own species. [12] Females duet with the males to help the male locate the females. [13] [14] Substrate-borne vibrations are not unique to treehoppers, most insects (more than 90%) use substrate-borne vibrations to communicate within species and between species. [13] There is a variety of ways insects can produce vibrations to communicate and even more variation in how they utilize vibrational communication (i.e. mating calls, cooperative foraging, or catching prey). [15]

Mating behavior and reproduction

Males search for mates by flying from one plant to another. As they land, they produce advertisement signals and wait for females to respond. [2] [11] [16] Different species in this clade are most divergent in the frequency (Hz) of their mating signals. [3] [17] [18] [19] [20]

Males fly or hop from plant to plant looking for female aggregations. [16] Females only mate once, while males mate multiples times. Soon after a female mates, she would start ovipositing eggs into the stem of the plant. Females have saw-like ovipositors that allow them to cut a slit into the plant stem and deposit her eggs. When the slits are filled up, she covers them with white secretions called egg froths. The egg froth protects the eggs from the elements, biotic and abiotic. [21]

Host-shifts and sympatric speciation

Assortative mating

Adult females mostly respond to conspecific signals. [22] [23] Females that are from a different host plant than the male rarely responds to the male's signals. If they do respond, there is even a lower chance of mating success.

Females have mating windows that conspecific males follow. This sort of reproductive isolation has contributed to the divergence of the clade. Males and females closer in age are more likely to form pre-copulatory and copulatory pairs. Larger time gaps between plant phenologies creates more disruption in the gene flow between sympatrically occurring species. [24]

Females tend to stay in their natal plants and prefer to mate and lay their eggs on it, which is called philopatry. [25] Eggs that were laid non-host plants have higher mortality due to different plant nutrition and the absence of native ants that nurture and protect nymphs. [11]

Phenology

Life histories of this species vary according to the phenology of their host plants. [21] These treehoppers lay their eggs on its host plant's branches, as well as spend their juvenile and adult life on one plant. [2] Egg hatching of these treehoppers are tied into the sap flow of their host plants. After winter, flow of the plant's sap to their stems is the stimuli the eggs need to start hatching. Once they have hatched from the stems as nymphs, they molt until adulthood (final form). Males start signaling first a week after they reach adulthood. Females become reproductively receptive 1–2 weeks about the males. After reproducing, females stay on one plant and oviposit their eggs continuously until they expire or until the first frost hits. Males live shorter than females and usually die shortly after mating a number of times.

Proposed phylogenetic tree for Enchenopa Proposed phylogenetic tree for Enchenopa.png
Proposed phylogenetic tree for Enchenopa

The complex has multiple species under the same name followed by the genus of its host plant (i.e. Enchenopa binotata 'Ptelea' for species that live in Ptelea trifoliata). [6] The nutritional value of the host plant's sap could delay or boost adult maturation and egg hatching. Sap with more essential nutrients or more sap flow for the species can promote faster maturation and/or egg hatching. [11] Species on different host plants have developed allochronic phenologies. This means that species on different host plants have evolved different timing in their life history. [21]

Male signals of the different species within the Enchenopa binotata complex Varying male signals that species of Enchenop binotata make.png
Male signals of the different species within the Enchenopa binotata complex

Phylogenetics

Phylogenetic data have showed that this species of Eastern North American treehoppers (E. binotata) diverged from two closely related species of Enchenopa from Central and South America. [27] These two Enchenopa species are known to be polyphagous (feeds on multiple species of plants) as opposed to E. binotata that are monophagous (only feeds on one species of plants). [3] [6] [11]

Male signal frequency is more heritable than female preference for certain frequencies. This means that there is more selection pressure on male signal frequency than female preference. Female choice might actually be the selection pressure that drove the divergence of male signal frequency in the E. binotata species complex. [3]

Host plants of species in the complex

Related Research Articles

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Hemiptera is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from 1 mm (0.04 in) to around 15 cm (6 in), and share a common arrangement of piercing-sucking mouthparts. The name "true bugs" is often limited to the suborder Heteroptera.

<span class="mw-page-title-main">Gerridae</span> Family of true bugs

The Gerridae are a family of insects in the order Hemiptera, commonly known as water striders, water skeeters, water scooters, water bugs, pond skaters, water skippers, water gliders, water skimmers or puddle flies. Consistent with the classification of the Gerridae as true bugs, gerrids have mouthparts evolved for piercing and sucking, and distinguish themselves by having the unusual ability to walk on water, making them pleuston (surface-living) animals. They are anatomically built to transfer their weight to be able to run on top of the water's surface. As a result, one could likely find water striders present in any pond, river, or lake. Over 1,700 species of gerrids have been described, 10% of them being marine.

<span class="mw-page-title-main">Leafhopper</span> Family of insects

Leafhopper is the common name for any species from the family Cicadellidae. These minute insects, colloquially known as hoppers, are plant feeders that suck plant sap from grass, shrubs, or trees. Their hind legs are modified for jumping, and are covered with hairs that facilitate the spreading of a secretion over their bodies that acts as a water repellent and carrier of pheromones. They undergo a partial metamorphosis, and have various host associations, varying from very generalized to very specific. Some species have a cosmopolitan distribution, or occur throughout the temperate and tropical regions. Some are pests or vectors of plant viruses and phytoplasmas. The family is distributed all over the world, and constitutes the second-largest hemipteran family, with at least 20,000 described species.

<span class="mw-page-title-main">Treehopper</span> Family of insects

Treehoppers and thorn bugs are members of the family Membracidae, a group of insects related to the cicadas and the leafhoppers. About 3,200 species of treehoppers in over 400 genera are known. They are found on all continents except Antarctica; only five species are known from Europe. Individual treehoppers usually live for only a few months.

<span class="mw-page-title-main">Auchenorrhyncha</span> Suborder of insects

The Auchenorrhyncha suborder of the Hemiptera contains most of the familiar members of what was called the "Homoptera" – groups such as cicadas, leafhoppers, treehoppers, planthoppers, and spittlebugs. The aphids and scale insects are the other well-known "Homoptera", and they are in the suborder Sternorrhyncha.

<i>Umbonia crassicornis</i> Species of true bug

Umbonia crassicornis, commonly known as the thorn bug, is a widespread member of the insect family Membracidae, and an occasional pest of ornamentals and fruit trees in southern Florida. The body length of the adult is approximately 10 millimetres (0.39 in). This is a variable species as to size, color and structure, particularly the pronotal horn of males. This tall, essentially perpendicular thornlike pronotum discourages birds and other predators from eating it, if only by mistakenly confusing it with a thorn. Typically, the adult is green or yellow with reddish lines and brownish markings.

<i>Amphipsalta zelandica</i> Species of true bug

The chorus cicada, Amphipsalta zelandica, is the most common species of cicada in New Zealand, where it is endemic and found in most areas. They typically live in forests and areas with open bush, where their left-over nymph skins can be seen on tree trunks and branches during the summer months. The males produce their cicada sound in unison, which can reach deafening proportions at the height of their population around February. Groups of cicada can suddenly transition from the typical cicada sound to synchronised clicks, using their wings to clap against the surface they are sitting on.

<span class="mw-page-title-main">Buffalo treehopper</span> Species of true bug

The buffalo treehopper is a species of treehopper belonging to the subfamily Smiliinae. It is sometimes classified as Ceresa bisonia.

<span class="mw-page-title-main">Seismic communication</span>

Seismic or vibrational communication is a process of conveying information through mechanical (seismic) vibrations of the substrate. The substrate may be the earth, a plant stem or leaf, the surface of a body of water, a spider's web, a honeycomb, or any of the myriad types of soil substrates. Seismic cues are generally conveyed by surface Rayleigh or bending waves generated through vibrations on the substrate, or acoustical waves that couple with the substrate. Vibrational communication is an ancient sensory modality and it is widespread in the animal kingdom where it has evolved several times independently. It has been reported in mammals, birds, reptiles, amphibians, insects, arachnids, crustaceans and nematode worms. Vibrations and other communication channels are not necessarily mutually exclusive, but can be used in multi-modal communication.

<span class="mw-page-title-main">Eriococcidae</span> Family of true bugs

Eriococcidae is a family of scale insects in the order Hemiptera. They are commonly known as felt scales or eriococcids. As of 2023, there are 109 genera and 681 species. Each species is usually specific to a different plant host, or closely related group of hosts.

<i>Centrotus cornutus</i> Species of true bug

Centrotus cornutus (thorn-hopper) is a species of "treehoppers" belonging to the family Membracidae.

<i>Aconophora compressa</i> Species of true bug

Aconophora compressa is a species of insect in the treehopper family, Membracidae. It is known by the common names lantana bug, lantana treehopper, lantana stemsucking treehopper, and lantana sap-sucking bug.

<i>Umbonia spinosa</i> Species of treehopper


Umbonia spinosa is a species of treehopper native to South America. It belongs to the Membracidae family.

<span class="mw-page-title-main">Aetalionidae</span> Family of true bugs

Aetalionidae are a family of treehoppers in the superfamily Membracoidea. Aetalionidae are somewhat like Membracidae in that they have one to three rows of short spines on the hind tibia but differ in having the front femur fused to the trochanter and the scutellum is completely exposed. The females have finger-like protrusions on the genital capsule. The family is mostly Neotropical. The subfamily Biturritiinae is Neotropical while the subfamily Aetalioninae has a Neotropical genus Aetalion and the sole Old World representative genus Darthula with a single species Darthula hardwickii.

<i>Entylia carinata</i> Species of true bug

Entylia carinata, commonly known as the keeled treehopper, is a species of treehopper in the family Membracidae. They can be found in Brazil, Panama, Mexico, the United States, and Canada. Keeled treehoppers are often attended by ants which feed on the honeydew they excrete. In return, the ants offer protection from predators. Keeled treehoppers typically feed on plants in the aster family and they are not known to transmit plant diseases and are not considered significant plant pests.

<i>Tylopelta gibbera</i> Species of true bug

Tylopelta gibbera is a species of treehopper in the family Membracidae. This insect has a range which extends from Guatemala into the central United States, although it is occasionally found in Canada. Vibrational communication has been noted in this species, although there is discussion as to what exactly it is being used for.

<i>Bocydium globulare</i> Species of true bug

The Brazilian treehopper is a species of insect belonging to the treehopper family (Membracidae). It has unusual appendages on its thorax. While Bocydium can be found throughout the world, they are most prevalent in Africa, North and South America, Asia and Australia. They exhibit limited movement and their primary food source is from the underside of leaves. They also exhibit hemimetabolous development. Treehoppers range from about two millimeters to two centimeters in length.

Laura Sullivan-Beckers is an associate professor of evolutionary biology at Murray State University. She is credited with the discovery of Hebetica sylviae, a species of treehopper, named for her daughter Sylvie Beckers.

<i>Enchenopa</i> Genus of true bugs

Enchenopa is a genus of treehoppers in the family Membracidae. There are more than 50 described species in Enchenopa.

<i>Cladonota</i> Genus of treehopper

Cladonota is a genus of neotropical treehoppers widespread from Mexico to South America. They are known for their elaborate pronotum shapes, hypothesized to play a role in camouflage or mate recognition, although their function is not yet known with certainty. Previously described as a subgenus of Sphongophorus, it was recognized as a genus of its own right in 1997 after the latter was synonymized with Hypsauchenia.

References

  1. ↑ [ dead link ]
  2. 1 2 3 4 Wood, T.K., 1980. Intraspecific divergence in Enchenopa binotata Say (Homoptera: Membracidae) effected by host plant adaptation. Evolution, 34: 147-160.
  3. 1 2 3 4 5 Fowler-Finn, K.D. et al. 2015. Variation in signal-preference genetic correlations in Enchenopa treehoppers (Hemiptera: Membracidae). Ecology and Evolution. 5(14): 2774-2786
  4. ↑ Deitz, L. L., & Wallace, M. S. (2012). Richness of the Nearctic treehopper fauna (Hemiptera: Aetalionidae and Membracidae). Zootaxa, 3423, 1-26.
  5. ↑ "Discover Life". Discoverlife.org. Archived from the original on 2016-09-18. Retrieved 2016-02-23.
  6. 1 2 3 4 5 6 7 8 9 10 11 12 13 Wood, Thomas K. (1993). "Speciation of the Enchenopa binotata complex (Insecta:Homoptera:Membracidae)". Evolutionary Patterns and Processes.
  7. ↑ [ dead link ]
  8. ↑ "Hemiptera: bugs, aphids and cicadas". Commonwealth Scientific and Industrial Research Organisation.
  9. ↑ "Photographic image of nymph progression" (JPG). Americaninsects.net. Archived from the original on 2017-01-18. Retrieved 2022-04-01.
  10. ↑ Cocroft, R. B., & McNett, G. D. (2006). 23 Vibratory Communication in Treehoppers (Hemiptera: Membracidae).
  11. 1 2 3 4 5 Wood, T.K. & Guttman, S.I., 1982. Ecological and behavioural basis for reproductive isolation in the sympatric Enchenopa binotata complex (Homoptera: Membracidae). Evolution, 36: 233-242.
  12. 1 2 Jocson, Dowen Mae I.; Smeester, Morgan E.; Leith, Noah T.; Macchiano, Anthony; Fowler-Finn, Kasey D. (2019-07-30). "Temperature coupling of mate attraction signals and female mate preferences in four populations of Enchenopa treehopper (Hemiptera: Membracidae)". Journal of Evolutionary Biology. 32 (10): 1046–1056. doi: 10.1111/jeb.13506 . ISSN   1010-061X. PMID   31278803.
  13. 1 2 Cocroft, Reginald B., and Rafael L. Rodríguez. (2005) "The behavioral ecology of insect vibrational communication." Bioscience 55.4: 323-334.
  14. ↑ Cocroft, R. B., Shugart, H. J., Konrad, K. T., & Tibbs, K. (2006). Variation in plant substrates and its consequences for insect vibrational communication. Ethology, 112(8), 779-789.
  15. ↑ Cocroft, R. B. (2005). Vibrational communication facilitates cooperative foraging in a phloem-feeding insect. Proceedings of the Royal Society of London B: Biological Sciences, 272(1567), 1023-1029.
  16. 1 2 Fowler-Finn, Kasey D., et al. "Male Enchenopa treehoppers (Hemiptera: Membracidae) vary mate-searching behavior but not signaling behavior in response to spider silk." Naturwissenschaften 101.3 (2014): 211-220.
  17. ↑ Mcnett, G.D. and Cocroft, R.B. (2008). Host shifts favor vibrational signal divergence in Enchenopa binotata treehoppers. Behavioral Ecology. 19.3: 650-656.
  18. ↑ Fowler- Finn, K.D., Rodriguez, R.L. 2012. Experience-mediated plasticity in mate preferences: mating assurance in a variable environment. Evolution. 66:459-468.
  19. ↑ Fowler-Finn, K.D., Rodriguez, R.L. 2012. The evolution of experience-mediated plasticity in mate preferences. Journal of Evolutionary Biology, 25(9): 1855-1863.
  20. ↑ Fowler-Finn, K.D., Rodriguez, R.L. 2013. Repeatability of mate preference functions in Enchenopa treehoppers (Hemiptera: Membracidae). Animal Behaviour. 1-7.
  21. 1 2 3 4 5 6 7 8 9 10 Wood, T. K. "Divergence in the Enchenopa binotata Say complex (Homoptera: Membracidae) effected by host plant adaptation." Evolution 34.1 (1980): 147-160.
  22. ↑ Hunt, R. E. (1994). Vibrational signals associated with mating behavior in the treehopper, Enchenopa binotata Say (Hemiptera: Homoptera: Membracidae). Journal of the New York Entomological Society, 102(2), 266-270.
  23. ↑ Rodríguez, Rafael L., Laura E. Sullivan, and Reginald B. Cocroft. "Vibrational communication and reproductive isolation in the Enchenopa binotata species complex of treehoppers (Hemiptera: Membracidae)." Evolution 58.3 (2004): 571-578.
  24. ↑ Wood, T. K., & Keese, M. C. (1990). Host-plant-induced assortative mating in Enchenopa treehoppers. Evolution, 619-628.
  25. ↑ Stearns, F. W., Tilmon, K. J., & Wood, T. K. (2013). Felsenstein's "one-allele model" of speciation: The role of philopatry in the initial stages of host plant mediated reproductive isolation in Enchenopa binotata. Current Zoology, 59(5), 658-666.
  26. 1 2 3 Cocroft, R. B., Rodríguez, R. L., & Hunt, R. E. (2008). Host shifts, the evolution of communication, and speciation in the Enchenopa binotata species complex of treehoppers. Specialization, speciation, and radiation: the evolutionary biology of herbivorous insects, 88-100.
  27. ↑ Guttman, Sheldon I., Thomas K. Wood, and Alvan A. Karlin. "Genetic differentiation along host plant lines in the sympatric Enchenopa binotata Say complex (Homoptera: Membracidae)." Evolution (1981): 205-217.
  28. 1 2 3 4 Rodriguez, R. L., Ramaswamy, K., and Cocroft, R. B. (2006). Evidence that females preferences have shaped male signal evolution in a clade of specialized plant-feeding insects. Proc. R. Soc. 273: 2585-2593.
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