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Tettigoniidae
Tettigoniidae
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Tettigoniidae
Temporal range: Jurassic–recent
Tettigonia viridissima
Stridulation of T. viridissima
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Orthoptera
Suborder: Ensifera
Infraorder: Tettigoniidea
Superfamily: Tettigonioidea
Krauss, 1902
Family: Tettigoniidae
Krauss, 1902
Subfamilies

See text

Insects in the family Tettigoniidae are commonly called katydids (especially in North America)[1] or bush crickets.[2] They have previously been known as "long-horned grasshoppers".[3] More than 8,000 species are known.[1] Part of the suborder Ensifera, the Tettigoniidae are the only extant (living) family in the superfamily Tettigonioidea.

Many species are nocturnal in habit, having strident mating calls and may exhibit mimicry or camouflage, commonly with shapes and colours similar to leaves.[4]

Etymology

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The family name Tettigoniidae is derived from the genus Tettigonia, of which the great green bush cricket is the type species; it was first described by Carl Linnaeus in 1758. In Latin tettigonia means a kind of small cicada, leafhopper;[5] it is from the Greek τεττιγόνιον tettigonion, the diminutive of the imitative (onomatopoeic) τέττιξ, tettix, cicada.[6][7] All of these names such as tettix with repeated sounds are onomatopoeic, imitating the stridulation of these insects.[8] The common name katydid is also onomatopoeic and comes from the particularly loud, three-pulsed song, often rendered "ka-ty-did", of the nominate subspecies of the North American Pterophylla camellifolia, belonging to the subfamily Pseudophyllinae, which are known as "true katydids".[9][10]

Description and life cycle

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Description

[edit]
Tettigonia viridissima

Tettigoniids range in size from as small as 5 mm (14 in) to as large as 130 mm (5 in).[11] The smaller species typically live in drier or more stressful habitats which may lead to their small size. The small size is associated with greater agility, faster development, and lower nutritional needs. Tettigoniids are tree-living insects that are most commonly heard at night during summer and early fall.[12] Tettigoniids may be distinguished from the grasshopper by the length of their filamentous antennae, which may exceed their own body length, while grasshoppers' antennae are always relatively short and thickened.

Katydid camouflaged on a bamboo leaf

Life cycle

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Katydid eggs attached in rows to a plant stem
Katydid nymph

Eggs are typically oval and may be attached in rows to plants. Where the eggs are deposited relates to the way the ovipositor is formed. It consists of up to three pairs of appendages formed to transmit the egg, to make a place for it, and place it properly. Tettigoniids have either sickle-shaped ovipositors which typically lay eggs in dead or living plant matter, or uniform long ovipositors which lay eggs in grass stems. When tettigoniids hatch, the nymphs often look like small, wingless versions of the adults, but in some species, the nymphs look nothing at all like the adult and rather mimic other species such as ants, spiders and assassin bugs, or flowers, to prevent predation. The nymphs remain in a mimic state only until they are large enough to escape predation. Once they complete their last molt (after about 5 successful molts), they are then prepared to mate.[12]

Distribution

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Tettigoniids are found on every continent except Antarctica.[13] The vast majority of katydid species live in the tropical regions of the world.[4] For example, the Amazon basin tropical forests are home to over 2,000 species.[4] However, katydids are found in the cool, dry temperate regions, as well, with about 255 species in North America.

Classification

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The Tettigoniidae are a large family and have been divided into a number of subfamilies:[1]

The Copiphorinae were previously considered a subfamily, but are now placed as tribe Copiphorini in the subfamily Conocephalinae.[14] The genus Acridoxena is now placed in the tribe Acridoxenini of the Mecopodinae (previously its own subfamily, Acridoxeninae).

Extinct taxa

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The Orthoptera species file[1] lists:

Genera incertae sedis
  • Locustites Heer, 1849: 3 spp.
  • Locustophanes Handlirsch, 1939: †L. rhipidophorus Handlirsch, 1939
  • Prophasgonura Piton, 1940: †P. lineatocollis Piton, 1940
  • Protempusa Piton, 1940: †P. incerta Piton, 1940
  • Prototettix Giebel, 1856: †P. lithanthraca (Goldenberg, 1854)

The genus †Triassophyllum is extinct and may be placed here or in the Archaeorthoptera.[15]

Ecology

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Poecilimon thoracicus (Phaneropterinae)

The diet of most tettigoniids includes leaves, flowers, bark, and seeds, but many species are exclusively predatory, feeding on other insects, snails, or even small vertebrates such as snakes and lizards. Some are also considered pests by commercial crop growers and are sprayed to limit growth, but population densities are usually low, so a large economic impact is rare.[16]

Tettigoniids are serious insect pests of karuka (Pandanus julianettii).[17] The species Segestes gracilis and Segestidea montana eat the leaves and can sometimes kill trees.[17] Growers will stuff leaves and grass in between the leaves of the crown to keep insects out.[17]

By observing the head and mouthparts, where differences can be seen in relation to function, it is possible to determine what type of food the tettigoniids consume. Large tettigoniids can inflict a painful bite or pinch if handled, but seldom break the skin.

Some species of bush crickets are consumed by people, such as the nsenene (Ruspolia differens) in Uganda and neighbouring areas.

Communication

[edit]

The males of tettigoniids have sound-producing organs located on the hind angles of their front wings. In some species, females are also capable of stridulation. Females chirp in response to the shrill of the males. The males use this sound for courtship, which occurs late in the summer.[18] The sound is produced by rubbing two parts of their bodies together, called stridulation. In many cases this is done with the wings, but not exclusively. One body part bears a file or comb with ridges; the other has the plectrum, which runs over the ridges to produce a vibration.[19] For tettigoniids, the fore wings are used to sing. Tettigoniids produce continuous songs known as trills. The size of the insect, the spacing of the ridges, and the width of the scraper all influence what sound is made.[20]

Many species stridulate at a tempo which is governed by ambient temperature, so that the number of chirps in a defined period of time can produce a fairly accurate temperature reading. For American katydids, the formula is generally given as the number of chirps in 15 seconds plus 37 to give the temperature in degrees Fahrenheit.[21]

Predation

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Wandering spider (Cupiennius sp.) with Tettigoniidae sp. prey

Some tettigoniids have spines on different parts of their bodies that work in different ways. The Listroscelinae have limb spines on the ventral surfaces of their bodies. This works in a way to confine their prey to make a temporary cage above their mouthparts. The spines are articulated and comparatively flexible, but relatively blunt. Due to this, they are used to cage and not penetrate the prey's body. Spines on the tibiae and the femora are usually more sharp and nonarticulated. They are designed more for penetration or help in the defensive mechanism they might have. This usually works with their diurnal roosting posture to maximize defense and prevent predators from going for their head.[22]

Defense mechanisms

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Katydid mimicking a leaf
A meadow katydid in Hawaii

When tettigoniids go to rest during the day, they enter a diurnal roosting posture to maximize their cryptic qualities. This position fools predators into thinking the katydid is either dead or just a leaf on the plant. Various tettigoniids have bright coloration and black apical spots on the inner surfaces of the tegmina, and brightly colored hind wings. By flicking their wings open when disturbed, they use the coloration to fool predators into thinking the spots are eyes. This, in combination with their coloration mimicking leaves, allows them to blend in with their surroundings, but also makes predators unsure which side is the front and which side is the back.[23]

 
Katydid

I LOVE to hear thine earnest voice,
   Wherever thou art hid,
Thou testy little dogmatist,
   Thou pretty Katydid!
Thou mindest me of gentlefolks, -
   Old gentlefolks are they, -
Thou say'st an undisputed thing
   In such a solemn way.

Thou art a female, Katydid!
   I know it by the trill
That quivers through thy piercing notes,
   So petulant and shrill.
I think there is a knot of you
   Beneath the hollow tree, -
A knot of spinster Katydids, -
   Do Katydids drink tea?

O, tell me where did Katy live,
   And what did Katy do?
And was she very fair and young,
   And yet so wicked, too?
Did Katy love a naughty man,
   Or kiss more cheeks than one?
I warrant Katy did no more
   Than many a Kate has done.

From the "To An Insect" poem by Oliver Wendell Holmes[24][25]

 
To A Katydid

LITTLE friend among the tree-tops,
       Chanting low your vesper hymns,
               Never tiring,
               Me inspiring,
       Seated 'neath the swaying limbs,
Do you know your plaintive calling,
When the summer dew is falling,
Echoes sweeter through my brain
Than any soft, harmonic strain?

Others call you an intruder,
       Say discordant notes you know;
               Or that sadness,
               More than gladness,
       From your little heart doth flow;
And that you awake from sleeping
Thoughts in quiet they were keeping,
Faithless love, or ill-laid schemes,
Hopes unanchored — broken dreams.

No such phantoms to my vision
       Doth your lullaby impart,
               But sweet faces,
               No tear traces,
       Smile as joyous in my heart,
As when first at mother's knee
Learned I your sweet mystery.
I defend you with my praises,
For your song my soul upraises.

Oft I fancy when your neighbors,
       In some secret thicket hid,
               Are debating,
               Underrating
       What that little maiden did,
That above their clam'rous singing
I can hear your accents ringing,
Like a voice that must defend
From abuse some time-loved friend.

Dream I not of fame or fortune,
       Only this I inward crave,
               Sweet assurance,
               Long endurance,
       Of a love beyond the grave.
Should my songs die out and perish,
You'll my name repeat and cherish;
Though all trace is lost of me,
Still you'll call from tree to tree.

From the "To A Katydid" poem by Kate Slaughter McKinney[26]

Reproductive behavior

[edit]

The males provide a nuptial gift for the females in the form of a spermatophylax, a body attached to the males' spermatophore and consumed by the female, to distract her from eating the male's spermatophore and thereby increase his paternity.[27]

Polygamy

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The Tettigoniidae have polygamous relationships. The first male to mate is guaranteed an extremely high confidence of paternity when a second male couples at the termination of female sexual refractoriness. The nutrients that the offspring ultimately receive will increase their fitness. The second male to mate with the female at the termination of her refractory period is usually cuckolded.[28]

Competition

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The polygamous relationships of the Tettigoniidae lead to high levels of male-male competition. Male competition is caused by the decreased availability of males able to supply nutritious spermaphylanges to the females. Females produce more eggs on a high-quality diet; thus, the female looks for healthier males with a more nutritious spermatophylax. Females use the sound created by the male to judge his fitness. The louder and more fluent the trill, the higher the fitness of the male.[29]

Stress response

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In species which produce larger food gifts, the female often seeks out the males to copulate. This, however, is a cost to females as they risk predation while searching for males. Also, a cost-benefit tradeoff exists in the size of the spermatophore which the male tettigoniids produce. When males possess a large spermatophore, they benefit by being more highly selected for by females, but they are only able to mate one to two times during their lifetimes. Inversely, male Tettigoniidae with smaller spermatophores have the benefit of being able to mate two to three times per night, but have lower chances of being selected by females. Even in times of nutritional stress, male Tettigoniidae continue to invest nutrients within their spermatophores. In some species, the cost of creating the spermatophore is low, but even in those which it is not low, it is still not beneficial to reduce the quality of the spermatophore, as it would lead to lower reproductive selection and success. This low reproductive success is attributed to some Tettigoniidae species in which the spermatophylax that the female receives as a food gift from the male during copulation increases the reproductive output of the reproduction attempt. However, in other cases, the female receives few, if any, benefits.[30]

The reproductive behavior of bush crickets has been studied in great depth. Studies found that the tuberous bush cricket (Platycleis affinis) has the largest testes in proportion to body mass of any animal recorded. They account for 14% of the insect's body mass and are thought to enable a fast remating rate.[31]

See also

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References

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[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
Tettigoniidae is a large and diverse family of in the order and suborder Ensifera, commonly known as katydids, long-horned grasshoppers, or bush crickets, also known in Brazilian Portuguese as esperanças (particularly in regions such as Espírito Santo), and sometimes as catadores or tucurás. With over 8,300 described species in approximately 1,300 genera across 19 subfamilies, it represents the largest family within Ensifera and is renowned for its members' acoustic signaling and leaf-like . Tettigoniids are distributed worldwide on all continents except , achieving their highest diversity in tropical regions where they occupy a variety of habitats including forests, grasslands, and shrublands. They exhibit filiform antennae longer than the body length, which serve for sensory detection, and forewings (tegmina) in many that mimic leaves, twigs, or bark to evade predators through or . Additional defenses include spines on the legs, chemical secretions, and a powerful bite in some taxa. Ecologically, most tettigoniids are herbivorous, feeding primarily on foliage, flowers, fruits, and seeds, though many species are omnivorous—incorporating pollen, fungi, and small invertebrates—or fully predatory on other insects and even snails. Primarily nocturnal and arboreal, they inhabit vegetation layers from understory to canopy, using stridulation (rubbing wings or legs together) to produce loud, species-specific songs for mate attraction and territorial defense, particularly by males. Females possess a prominent, sword-like ovipositor for inserting eggs singly or in clusters into plant tissues or soil, and nymphs undergo incomplete metamorphosis while resembling adults. Certain species can be agricultural pests by damaging crops, while others play roles as prey in food webs or indicators of habitat health.

Taxonomy and phylogeny

Classification

Tettigoniidae is classified within the order , suborder Ensifera, and superfamily Tettigonioidea, where it represents the sole extant family. This placement distinguishes it from other ensiferans like (Grylloidea) and (Gryllotalpoidea), emphasizing its unique position among long-horned orthopterans. As of 2025, Tettigoniidae encompasses 8,493 described species distributed across 1,353 genera, reflecting ongoing discoveries and taxonomic refinements from phylogenetic analyses. These estimates are maintained in authoritative databases like the Orthoptera Species File, which track the family's diversity amid rapid in tropical regions. Historically, the classification of Tettigoniidae has undergone significant revisions, particularly in the when many groups initially described as separate families by Brunner von Wattenwyl (1878) were later consolidated into subfamilies under Tettigoniidae. A nomenclatural review further standardized family-group names. More recent molecular phylogenies, such as those from 2018, have further adjusted boundaries, including the synonymization of paraphyletic groups like Meconematinae to better align with evolutionary relationships. Key diagnostic traits for identifying Tettigoniidae at the family level include antennae that are filiform and typically longer than the body length, often exceeding it by several times, and in females, a prominent, elongate that is usually sword-like or needle-shaped for egg deposition. These features, combined with the vertical orientation of the wings at rest, reliably separate tettigoniids from shorter-antennaed orthopterans like acridids.

Subfamilies and genera

The family Tettigoniidae encompasses 19 subfamilies, containing 8,493 described distributed across 1,353 genera as of 2025. These subfamilies exhibit significant morphological and ecological diversity, ranging from leaf-mimicking forms to predatory specialists, with the majority of species concentrated in tropical regions. Prominent subfamilies include Phaneropterinae, the largest with more than 2,100 of often leaf-like katydids found worldwide; Pseudophyllinae, comprising over 1,000 in about 240 genera, predominantly in tropical and subtropical habitats; Conocephalinae, with approximately 1,300 characterized by elongated heads in many taxa; Meconematinae, including around 900 of smaller, often predatory forms; and , featuring around 1,000 of shield-backed katydids mainly in temperate zones. Other significant subfamilies are , Listroscelidinae (spiny predatory katydids), Saginae, Hetrodinae, and Bradyporinae.
SubfamilyApproximate Species CountKey Characteristics and Distribution
Phaneropterinae>2,100Leaf-like ; global, especially . (approx. 2018)
Pseudophyllinae>1,000True katydids; tropical/subtropical focus. (approx. 2024)
Conocephalinae~1,300Cone-headed forms; cosmopolitan. (approx. 2018)
Meconematinae~900Small, predatory; emphasis. (approx. 2018)
Tettigoniinae~1,000Shield-backed; temperate regions. (approx. 2018)
Notable genera highlight regional endemism and adaptations, such as Mecopoda in , a group of large katydids distributed across tropics including , , and ; and Scudderia in Phaneropterinae, North American bush katydids renowned for leaf-mimicking wings and occurring in deciduous forests and shrublands. Recent taxonomic revisions, informed by phylogenetic studies, have reclassified the former subfamily Copiphorinae as the tribe Copiphorini within Conocephalinae, reflecting monophyletic groupings based on molecular data. Diversity hotspots are evident in the Neotropics, where over 2,000 Tettigoniidae species inhabit the , underscoring the region's role in katydid .

Evolutionary history

The family Tettigoniidae originated during the period, with molecular clock estimates placing the crown-group divergence around 155 million years ago based on fossil-calibrated phylogenomic analyses of mitochondrial and nuclear markers. More recent 2025 molecular clock studies using mitochondrial phylogenomics corroborate an origin in the , between approximately 150 and 200 million years ago, incorporating fossil calibrations from ensiferan outgroups. Key divergence events within Tettigoniidae occurred during the , with major subfamilies separating around 100 million years ago amid the breakup of and the diversification of early flowering plants. Following the Cretaceous-Paleogene (K-Pg) boundary approximately 66 million years ago, the family underwent significant , particularly in tropical regions, facilitated by the expansion of angiosperm-dominated forests that provided new ecological niches for and . Phylogenetically, Ensifera (including Tettigoniidae) forms a monophyletic to within , supported by comprehensive analyses of over 100 orthopteran species using multiple genetic loci. Tettigoniidae itself is strongly monophyletic, with robust support from efforts targeting the COI gene, as demonstrated in 2025 regional surveys that resolved familial boundaries across diverse taxa. These radiations are linked to the evolution of angiosperms, enabling leaf-mimicking in many lineages, alongside adaptations to nocturnal lifestyles that reduced competition and predation risks in forest understories.

Extinct taxa

Fossils of Tettigoniidae are documented from Permian to Eocene deposits, providing key insights into the family's early diversification. Major fossil sites include the Middle Permian of , the Late Jurassic Karatau Formation in , and Eocene oil shales in , such as the Green River Formation in . Approximately 20 extinct genera have been described within Tettigoniidae, highlighting a rich paleodiversity. Notable examples include Permotettigonia gallica from the Middle Permian of , an early stem-group exhibiting leaf-mimicking wing morphology as an against predation. Another is Aboilus aulietus from the of , characterized by distinctive forewing venation patterns. The genus Pseudotettigoniinae encompasses several species, with fossils showing transitional features between ancient and modern forms. These extinct taxa reveal evolutionary significance in the development of acoustic communication. Early forms display primitive stridulatory organs, such as the file-and-scraper mechanism on the forewings of Archaboilus musicus from the of , which produced low-frequency musical calls for mate attraction. evidence also confirms origins for certain subfamilies, like the Simoderinae, with relictual distributions in southern continents suggesting vicariance following the breakup of . Recent discoveries in the 2020s from amber deposits in have uncovered Tettigoniidae inclusions preserving fine details of wing venation and potential acoustic structures, indicating advanced production capabilities by the mid-. Similarly, a 2023 Eocene fossil of Arethaea solterae from the Green River Formation exceptionally preserves internal organs, including muscles and glands, shedding light on soft-tissue in the family.

Morphology and development

Physical characteristics

Tettigoniidae, commonly known as katydids or long-horned grasshoppers, exhibit a robust adapted for life in , with a pronotum that is often enlarged and saddle-shaped, appearing shield-like in some subfamilies such as . The body is typically taller than wide and thin, frequently leaf-shaped to facilitate , with sizes ranging from small species under 10 mm to large ones with wingspans exceeding 250 mm. They possess chewing mouthparts and long, slender legs, including enlarged hind femora suited for jumping, while the front and middle legs may bear spines in predaceous species for grasping prey. A defining feature is the filiform antennae, which are thread-like and typically as long as or longer than the body, sometimes reaching 3-4 times its length, and equipped with numerous sensory receptors. The auditory organs, or tympana, are located on the front tibiae, aiding in sensory perception. Wing morphology varies widely, with the forewings (tegmina) often leathery and held roof-like over the when present; they serve functions including sound production via a stridulatory file on the male's forewings. Many species are fully winged, but others are brachypterous with reduced tegmina or apterous, lacking functional wings altogether, which limits flight capability in those forms. Sexual dimorphism is pronounced, particularly in the abdomen: females are generally larger and bear a long, sword-like or sickle-shaped ovipositor for egg-laying, which can be flattened and saber-like or curved depending on the species. Males, in contrast, have short, non-articulated cerci used in spermatophore transfer, along with specialized wing structures for stridulation. Coloration in Tettigoniidae is predominantly or to blend with foliage or bark, with many species displaying leaf-like patterns or of twigs and dried for concealment; rare variants may exhibit , appearing pink.

Life cycle stages

Tettigoniidae, commonly known as katydids or bush crickets, undergo hemimetabolous or incomplete , characterized by three primary life stages: , , and , without a distinct pupal phase. In this developmental pattern, resemble miniature but lack fully developed wings and reproductive organs, gradually acquiring these features through a series of molts. The process ensures that young stages are adapted for growth and dispersal, with wing pads appearing in later instars to facilitate eventual flight in . The stage represents the initial and often overwintering phase for many . Females typically deposit in clusters within , plant stems, or slits cut into vegetation using their , with being the ancestral and common site for numerous taxa. In temperate , enter —a dormant state triggered by environmental cues like shortening day length—to survive winter, in spring after 6-9 months of development; tropical , by contrast, exhibit direct development without , allowing multiple generations annually. size and number vary by , but deposition often occurs in late summer or fall to align with favorable conditions. Nymphs emerge from eggs as first-instar juveniles and progress through 4-8 molts, depending on species, nutrition, and temperature, with each instar lasting about 1-2 weeks. The total nymphal period spans 1-3 months, during which they feed voraciously on foliage and undergo ecdysis to shed exoskeletons, revealing progressively larger body sizes, more defined wing pads, and enhanced camouflage patterns that mimic leaves or twigs for predator avoidance. Early instars are highly vulnerable and gregarious in some species, while later ones become more solitary and mobile. Upon completing the final molt, nymphs emerge as sexually mature adults, with full wing development enabling stridulation and flight. Adult lifespans range from 1-6 months, influenced by , predation, and resource availability, during which they focus on before . is attained immediately post-molt, allowing rapid pairing and oviposition to perpetuate the cycle.

Distribution and habitat

Global distribution

Tettigoniidae exhibit a , with present on all continents except . The family comprises over 8,400 valid extant , the vast majority of which occur in tropical and subtropical regions, reflecting their preference for warmer climates. Diversity is notably low in polar areas and extreme arid zones, where environmental conditions limit their establishment. Regional patterns of diversity highlight the Neotropics as the primary hotspot, harboring approximately 1,800 described species across diverse subfamilies, though no endemic subfamilies are present. The Oriental region supports high species richness, particularly in , with significant contributions from subfamilies such as Phaneropterinae (over 2,600 species globally, many Asian) and Meconematinae. follows with around 1,350 species, while other regions like and Afrotropics also show substantial but lower counts. The family's biogeographic history traces back to ancient Gondwanan origins in the , approximately 155 million years ago, followed by diversification across southern continents and multiple transoceanic dispersals that facilitated their global spread. More recently, human activities have aided introductions, such as the expansion of Conocephalus discolor across since the late 20th century. Advances in molecular techniques, including , have revealed substantial cryptic diversity in recent years, particularly in and , leading to upward revisions in species counts. For instance, integrative taxonomic studies in West African rainforests have uncovered hidden lineages within bush-cricket genera, while analyses in have identified species complexes previously unrecognized.

Habitat preferences

Tettigoniidae species occupy a diverse array of primary habitats worldwide, including forests, grasslands, shrublands, savannas, deserts, and rocky mountain tops. In tropical regions, many species are arboreal, residing in the canopy and of dense , particularly within subfamilies like Phaneropterinae and Pseudophyllinae, which favor tall herbaceous and trees. In contrast, temperate zone species tend to be more ground-dwelling or associated with low vegetation in open areas, as seen in Tettigoniinae, which prefer Mediterranean shrublands and grasslands. These preferences reflect the family's broad adaptability to vegetated environments from coastal littorals to high-elevation zones. Microhabitat selection within these broader environments often involves concealed or structurally complex sites that provide and protection. Many utilize leaf , understory vegetation, and soil layers for resting and oviposition, with Conocephalinae found across canopy, , and strata in forests. Some taxa, such as members of Pseudophyllinae, inhabit margins of aquatic or marine environments, including rock crevices along littorals or near bodies, while others like Microtettigoniinae associate with grasses and lilies in herbaceous settings. These microhabitats enhance and reduce predation risk, aligning with the family's reliance on vegetative cover. The family exhibits a wide altitudinal range, from to over 4,000 meters in mountainous regions, with like those in Eupholidoptera occurring from coastal lowlands to 1,800 meters and others, such as the , extending above 3,300 meters in . Desert-adapted , including those in genera like Idiostatus and certain Thar Desert tettigoniids, show xeric modifications such as heat storage behaviors and mimicry to endure arid conditions with limited water and vegetation. These adaptations enable persistence in fragmented, stressful habitats like valleys and desert edges. Climate plays a pivotal role in shaping Tettigoniidae distributions and activity patterns, with highest in humid tropical and subtropical zones that support lush . Temperate species, particularly in , exhibit seasonal activity tied to warmer months, entering or reduced mobility during cold winters, which limits their presence in regions with prolonged low s. This climatic sensitivity underscores the family's dependence on moderate and for optimal utilization.

Ecology and interactions

Diet and foraging

Tettigoniidae exhibit an omnivorous diet, with most primarily herbivorous, consuming leaves, flowers, stems, fruits, and seeds from a diverse array of plants. Analysis of digestive tract contents from Neotropical katydids reveals no strong dietary specialization, as individuals feed on multiple plant families, including and , reflecting generalist herbivory across canopy and vegetation. However, many supplement their plant-based diet with animal matter, particularly smaller like and other arthropods, with some lineages, such as certain Phaneropterinae, showing predominantly predatory habits. Foraging in Tettigoniidae is predominantly nocturnal, allowing individuals to browse or prey under cover of darkness while minimizing exposure to diurnal predators. employ varied strategies, including slow browsing on foliage for material and stationary ambushes for , with some, like those in the Phaneroptera, specializing in and consumption from flowers. This flexibility enables to in diverse habitats. Nutritional ecology in Tettigoniidae involves opportunistic , especially among nymphs, which consume molting or injured conspecifics to acquire essential proteins and salts in nutrient-poor environments, as observed in species like the Anabrus simplex. In adults, males transfer substantial nutrients to females via spermatophylax nuptial gifts during mating, providing proteins that enhance female fecundity and egg production by up to 85% under food restriction. Certain Tettigoniidae achieve pest status due to their herbivory on crops; for instance, Segestidea montana defoliates (Pandanus julianettii) trees in Papua New Guinea's highlands, damaging this culturally and nutritionally vital nut crop.

Predation dynamics

Tettigoniidae, commonly known as katydids, serve as important prey in various ecosystems, particularly in tropical forests where they occupy a mid-level trophic position as both herbivores and occasional predators of smaller . Major predators include birds such as flycatchers (e.g., La Sagra's flycatcher, Myiarchus sagrae), which actively forage for katydids in vegetation; bats, notably gleaning species like Micronycteris hirsuta that detect prey through echolocation and incidental sounds; spiders, which ambush resting individuals; and , which consume them opportunistically on foliage. These interactions position katydids as a key biomass contributor in tropical canopies, where their abundance supports higher trophic levels, with studies indicating intermediate trophic positions relative to primary consumers like grasshoppers and top predators like spiders in systems. Anti-predator cues play a critical role in katydid survival, particularly against nocturnal threats. Katydids possess ultrasound-sensitive ears that allow acoustic detection of bat echolocation calls, triggering immediate escape responses such as song cessation or flight, which reduces predation risk from gleaning bats. Predation intensity often peaks seasonally, aligning with katydid calling periods at night when males are most vulnerable, though bats can also target silent females and nymphs. In food webs, katydids function as vital prey for insectivores, sustaining populations of birds, bats, and reptiles while influencing dynamics. Their outbreaks can elevate herbivory rates, potentially defoliating vegetation, but predation pressure from these consumers helps regulate katydid densities, thereby moderating impacts on communities in tropical and temperate habitats. This top-down control underscores their role in maintaining and carbon cycling in forest .

Defense strategies

Tettigoniidae employ a variety of defense strategies to evade predators, primarily relying on , aposematic displays, physical barriers, and behavioral responses. These adaptations are particularly crucial given their vulnerability to visually hunting arthropods, birds, and acoustically foraging bats. through leaf or twig is a primary defense in many Tettigoniidae, achieved via specialized body coloration, wing venation, and resting postures that blend with foliage. For instance, in the genus Phyllophora (Phyllophorinae) exhibit green or brown hues with irregular outlines and assume flattened postures on branches to resemble dead leaves, reducing detection by visual predators. Leaf-like wings have evolved multiple times across Tettigoniidae subfamilies, enhancing this by providing irregular, vein-patterned surfaces that disrupt body outlines. Chemical defenses occur in select subfamilies, often paired with startle displays to deter close-range attackers. In the Phaneropterinae, species like Vestria produce glandular secretions believed to act as repellents when disturbed. Similarly, the mountain katydid Acripeza reticulata (Acripezinae) regurgitates bitter crop fluids and releases abdominal secretions containing alkaloids, which taste unpalatable to predators such as birds and . These chemicals are frequently revealed alongside deimatic displays, where the flashes brightly colored hindwings—such as the blue and orange eyespots in Pterochroza ocellata (Phaneropterinae)—to startle or confuse assailants, buying time for escape. Physical defenses include morphological structures and reflexive behaviors suited to their arboreal or ground-dwelling habits. Many Tettigoniidae possess sharp spines on their legs and , as seen in genera like Steirodon (Steirodontinae), which can injure or deter grasping predators such as spiders and mantids. Thanatosis, or feigning death by remaining motionless in a rigid posture, is employed by some when threatened, mimicking inedible debris to discourage further investigation. serves as a rapid escape mechanism, powered by elongated hind legs that propel the distances up to several body lengths, often combined with flight in winged forms. Acoustic defenses target auditory predators, particularly , through both passive and active means. Numerous Neotropical Tettigoniidae reduce calling during periods of high bat activity, entering silent phases to avoid by bats that home in on mate-attraction songs. When directly threatened, some species produce defensive , such as the broadband hissing sounds generated by rubbing forewings in Poecilimon ornatus (Poecilimoninae), which may startle vertebrate predators or interfere with attack coordination. These ultrasonic or low-frequency emissions can also disrupt bat echolocation in close encounters, though cessation of signaling remains the predominant anti-bat strategy.

Behavior and communication

Acoustic signaling

Tettigoniidae, commonly known as katydids or bush crickets, produce sounds primarily through , a mechanism involving the rubbing of specialized structures on their forewings, or tegmina. In males, which are the primary sound producers, one tegmen features a file—a series of ridges or teeth—while the other has a scraper or that rubs against it during wing closure, generating vibrations amplified by the wing's resonant properties. This process creates species-specific songs that vary in pattern and frequency, often consisting of trills, chirps, or pulses. For instance, carrier frequencies typically range from 2 to 50 kHz, with some species producing low-frequency audio calls around 5-13 kHz and others extending into above 30 kHz, allowing adaptation to diverse acoustic environments. These songs are predominantly nocturnal, with stridulation typically occurring at night and often continuing into the early morning hours, producing persistent and repetitive calls that can be audible over prolonged periods. These acoustic signals serve multiple functions in Tettigoniidae communication, including mate attraction and territorial defense. Males broadcast calling songs to lure receptive females over long distances, with often acting as a recognition cue to prevent hybridization. In territorial contexts, aggressive songs or rival interactions help maintain spacing among males, reducing competition for resources and mates. Additionally, some emit ultrasonic signals, particularly in response to predation threats from echolocating bats, triggering avoidance behaviors such as flight cessation or directional steering away from the sound source. In tropical regions of Brazil, such as Espírito Santo, these persistent nocturnal sounds are sometimes perceived by local residents as annoying or bothersome ("barulho chato"), particularly during hot and humid periods when insect activity is heightened. Hearing in Tettigoniidae is mediated by tympanal organs located in the proximal of the forelegs, consisting of thin membranes that vibrate in response to airborne sounds and connect to sensory neurons. These organs exhibit frequency tuning, with sensitivity peaks aligned to the dominant frequencies of conspecific songs, enabling precise detection amid . The auditory system also responds to for predator detection, integrating dual roles in communication and survival. Recent bioacoustics research has highlighted habitat-specific variations in Tettigoniidae calls from the , where studies in 2025 revealed diverse song patterns among syntopic in the .

Social behaviors

Tettigoniidae exhibit a range of social behaviors, with most displaying solitary habits as adults, particularly in low-density conditions where individuals maintain spatial separation to minimize competition and predation risk. However, certain demonstrate density-dependent gregariousness during outbreaks, shifting to aggregated formations that facilitate movement and enhance . For instance, the (Anabrus simplex) transitions from a cryptic, sedentary solitary phase to a gregarious phase characterized by aposematic coloration, band formation, and migration, with densities exceeding solitary populations by over a thousandfold; these outbreaks trigger behaviors such as increased mobility and group foraging, observed through radiotelemetry showing distinct movement patterns in high-density versus low-density groups. Nymphal aggregation occurs in gregarious species like the , where early instars cluster into mobile bands for protection against predators and environmental stressors, potentially aiding regulation though direct thermoregulatory benefits remain understudied in Tettigoniidae. While pheromone-mediated trails are not widely documented in this , acoustic cues from choruses can promote aggregative spacing in singing males, as seen in bush-cricket assemblages where influences regular inter-male distances to optimize overlap. Territoriality is prominent among males, who defend calling sites through acoustic interactions that serve as contests for resource access, including oviposition and feeding areas in herbivorous species. In Tettigonia cantans, males assess using song carrier frequency (with dominants producing lower frequencies by approximately 1 kHz), body weight (heavier individuals more likely to dominate), and prior site occupation, leading to agonistic escalations that establish without physical contact in most cases. Similarly, in Mygalopsis marki, competing males modify call temporal patterns, such as duration and rate, to outcompete and retain territorial control. These behaviors ensure exclusive access to high-quality herbaceous patches, reducing interference in phytophagous groups. Interspecific interactions in Tettigoniidae often involve for predator avoidance, with some nymphs resembling other to exploit model species' defenses. Neotropical katydids frequently employ or Batesian strategies, including ant-like morphologies in early instars of certain Phaneropterinae, which deter attacks by mimicking unpalatable or defended ; leaf-mimicking adults in genera like Typophyllum may indirectly benefit from ant associations in shared arboreal habitats, though true remains rare and unconfirmed. These adaptations highlight non-reproductive social dynamics shaped by ecological pressures rather than intraspecific signaling.

Reproduction and mating

Mating systems

In Tettigoniidae, mating systems are characterized by male investment in nuptial gifts and acoustic attraction, with females often exercising choice based on male signals and offerings. Males typically produce species-specific songs to attract receptive females , a process rooted in acoustic signaling that facilitates initial mate location. Once a female approaches, rituals ensue, involving close-range behaviors such as antennal touching to assess compatibility and, in some species, wing fanning to display fitness or release pheromones. These interactions ensure mutual recognition and alignment before copulation, emphasizing sensory cues beyond long-range calls. A hallmark of tettigoniid reproduction is the transfer of a large during , which serves dual roles in delivery and nutritional provisioning to the female. The consists of a -containing and a gelatinous spermatophylax that the female consumes post-copulation, providing essential proteins and equivalent to up to 30% of the male's body weight in some species. This nuptial gift enhances female by supporting egg production, while also prolonging copulation to increase the chances of successful transfer by deterring immediate female removal of the . Many tettigoniid species exhibit promiscuous mating patterns, with prevalent as females mate multiply to acquire multiple nuptial gifts and diverse for potential genetic benefits. Female choice plays a central role, favoring males with superior song quality—indicating genetic or condition-based fitness—or larger spermatophores that signal resource investment. While some taxa show monogamous tendencies, dominates in resource-limited environments, driving through both pre- and post-copulatory mechanisms. Nuptial gifts vary across subfamilies, being prominent in but reduced or absent in others like Phaneropterinae. Mating activity in Tettigoniidae peaks nocturnally, aligning with the nocturnal calling patterns of males to minimize predation risk and optimize signal propagation in low-light conditions. In temperate regions, reproduction is seasonal, occurring primarily after nymphal emergence in late summer, with calling and mating intensifying from July through October as adults reach sexual maturity. This timing synchronizes with environmental cues like temperature and photoperiod to maximize reproductive success within the adult lifespan.

Reproductive competition

In Tettigoniidae, male-male for mates often involves physical confrontations, particularly in where males defend calling sites or females. Males in some conehead katydids engage in aggressive battles to establish dominance and secure opportunities. These fights can include grappling, biting, and targeting vulnerable areas such as the opponent's eyes, with larger or more robust males typically prevailing. Sperm competition is a prominent form of post-copulatory rivalry in polyandrous Tettigoniidae species, where females mate with multiple partners and store spermatophores from successive males in their . The spermatophore consists of a sperm-containing and a nutrient-rich spermatophylax, allowing females to retain ejaculates from several males, which compete to fertilize eggs. In many cases, last-male precedence dominates, with the most recent male siring up to 90% of , as observed in Poecilimon veluchianus, promoting strategies like removal or oversized spermatophores to displace prior rivals. Female-female competition in Tettigoniidae manifests in species with role-reversed mating systems, where females aggressively vie for access to males providing nutritious nuptial gifts. For instance, in Kawanaphila nartee, females compete directly for singing males by displacing rivals through physical aggression, leading to for enhanced auditory sensitivity via larger thoracic spiracles in competitively superior females. Although direct guarding of oviposition sites is less documented, aggressive interactions can indirectly influence resource access during egg-laying, as females with priority secure better nutritional states for subsequent oviposition. These competitive dynamics are driven by , favoring exaggerated male traits such as enlarged cerci used for grasping females during copulation or subduing rivals. In species like those in the , hyperallometric growth of cerci and other genital structures reflects intense selection pressures, enhancing mating success amid high and rival interference.

Physiological responses

In Tettigoniidae, stress responses during reproductive activities, such as or exposure to predation risks, involve physiological adjustments in composition to bolster immunity. Under nutritional stress or protein deficiency, protein levels decrease, reducing resistance to entomopathogens like in species such as the (Anabrus simplex), highlighting how resource limitation impairs stress tolerance. stress further modulates immune activation, with production trading off against immune function; in some bushcricket species, larger spermatophores correlate with reduced phenoloxidase activity, a key immune enzyme, indicating energy reallocation from immunity to reproduction. Post-copulatory physiological changes in females include a that renders them unreceptive to remating, often lasting several days to weeks and linked to alterations in reproductive such as delayed oviposition timing. In Requena verticalis, this period is induced by substances in the , promoting over remating intervals and enhancing female nutrition via nuptial gifts while limiting multiple matings. Males experience significant exhaustion from nuptial gift production, with the energy content of spermatophores representing up to 25% of a male's body energy reserves in like Kawanaphila nartee, constraining subsequent reproductive efforts and increasing vulnerability to stressors. Hormonal regulation plays a critical role in timing reproductive events under stress. influences oviposition and receptivity in Tettigoniidae, maintaining reproductive readiness while interacting with nutritional status to modulate egg production rates. supports stress-induced physiological adjustments in reproductive processes, as observed in related where levels fluctuate with environmental pressures. Pathogen resistance in reproductive contexts is enhanced by in and reproductive fluids, particularly in dense populations where disease transmission risks rise. In Anabrus simplex, density-dependent prophylaxis elevates immune responses, including activity, in crowded conditions to counter pathogens, a mechanism vital for migratory swarms. Recent studies on bush-crickets confirm trade-offs between reproductive investment and immunity, with proteins potentially activating defenses to protect reproductive tissues amid heightened disease exposure in high-density aggregations.

References

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