View source for Tingidae

The '''Tingidae''', commonly referred to as '''lace bugs''' due to their intricate wings, are a [[Family (biology)|family]] of very small (2-10 mm (0.08-0.39 in)) [[Insect|insects]] in the order [[Hemiptera]].<ref name=":05">{{Citation |last1=Guidoti |first1=Marcus |title=Lace Bugs (Tingidae) |date=2015 |work=True Bugs (Heteroptera) of the Neotropics |volume=2 |pages=395–419 |editor-last=Panizzi |editor-first=Antônio R. |url=https://link.springer.com/chapter/10.1007/978-94-017-9861-7_14 |access-date=2025-03-22 |place=Dordrecht |publisher=Springer Netherlands |language=en |doi=10.1007/978-94-017-9861-7_14 |isbn=978-94-017-9860-0 |last2=Montemayor |first2=Sara I. |last3=Guilbert |first3=Éric |editor2-last=Grazia |editor2-first=Jocélia|url-access=subscription }}</ref> These insects exist in multiple regions of the world and live on various plants, depending on the [[species]]. There are roughly 2,000 described species across the 3 [[Subfamily|subfamilies]] which include Cantacaderinae, Tinginae and Vianaidinae.{{Short description|Family of true bugs}}
{{Automatic taxobox
| name = Tingidae
| fossil_range = {{fossil range|Early Cretaceous|Present}}
| image = HEMI Tingidae Tanybyrsa cumberi.png
| image_caption = ''[[Tanybyrsa cumberi]]''
| taxon = Tingidae
| authority = [[Francis de Laporte de Castelnau|Laporte]], 1832
| display_parents = 2
| subdivision_ranks = Subfamilies
| subdivision = *{{extinct}}[[Tingiometrinae]] <small>Heiss, Golub, & Popov, 2015</small> 
*[[Cantacaderinae]] <small>Stål, 1873</small>
*[[Tinginae]] <small>Laporte, 1832</small>
*[[Vianaidinae]] <small>Kormilev, 1955</small>
}}

== Characteristics ==
They have two main life stages, nymphal and adult. The nymphal period consists of 5 [[instar]] stages each similar but unique in their own manners lasting a total of 8-16 days.<ref name=":10">{{Cite journal |last1=Kumar |first1=Nagendra |last2=Kumar |first2=Anil |date=2018 |title=Study the Biology of Lace Bug, Cochlochila Bullita (Stal) (Hemiptera: Tingidae) Under Ambient Laboratory Conditions |url=https://www.phytojournal.com/special-issue/2018.v7.i1S.3462/study-the-biology-of-lace-bug-cochlochila-bullita-stal-hemiptera-tingidae-under-ambient-laboratory-conditions |journal=Journal of Pharmacognosy and Phytochemistry |language=en |volume=7 |issue=1S |pages=1755–1760 |issn=2278-4136}}</ref> The first instar stage occurs after hatching and lasts typically 2-4 days, the molt is oblong shaped and transforms from a pale brown to a brownish-black coloured [[Cuticle (arthropod)|cuticle]]. The second instar also typically lasts 2-4 days, it remains a dark brown-black colour but the molt is rounded instead, the abdominal surface have indications of spinal projections along the body's margin. The third instar typically lasts 2-3 days and the molt remains round but larger in size with short cephalic spines. The fourth instar lasts 1-3 days and the molt continues to grow in size, large cephalic spines as well as a wider paranotum on the thorax are observed. Finally, the fifth instar stage lasts 1-2 days and is very similar to the previous stage but the molt transforms back into an elongated body form. This concludes the nymphal stage, transforming the cuticle into its adult life form.

The adult life forms are quite small with typical body size ranging anywhere from 2-10 mm (0.08-0.39 in),<ref name=":05" /> varying in shape, from quadrate, broadly oval or narrow. Tingidae are sexually differentiated with females being larger and having an [[ovipositor]] while the males are smaller with a genital capsule.<ref name=":10" /> They are cryptic in colouration, varying from the whitish-yellow range to dark brown and black. They have very intricate partially transparent hyaline forewings and pronotum wings, the detailing is lace-like giving Tingidae the common name "lace bug".<ref name=":9">{{Citation |last=Miller |first=Laura T. |title=Lace Bugs (Hemiptera: Tingidae) |date=2008 |encyclopedia=Encyclopedia of Entomology |pages=2099–2102 |editor-last=Capinera |editor-first=John L. |url=https://link.springer.com/referenceworkentry/10.1007/978-1-4020-6359-6_1934 |access-date=2025-03-25 |place=Dordrecht |publisher=Springer Netherlands |language=en |doi=10.1007/978-1-4020-6359-6_1934 |isbn=978-1-4020-6359-6|url-access=subscription }}</ref> On the [[Thorax (arthropod anatomy)|thorax]] is the pronotum as well as a hollow outgrowth on the dorso-lateral region. They have 2 sets of wings; a pair of forewings and pronotum wings, which vary in size between individuals, typically having reduced forewings and larger pronotum wings. Despite their apparent beauty, the wings are highly decorative and adults are poor fliers, carrying out their sedentary lifestyle on leaves and the shoots of plants.<ref name=":10" />

Due to their lifestyle, Tingidae require unique cephalic characteristics. Their heads are either fully concealed or partially obscured under a hood-like projection arising from the pronotum. Some develop spine-like projections while others do not, and all possess a 4-way segmented antennae, with the third one being the longest.<ref name=":9" /> The lack of [[Simple eye in invertebrates|ocelli]]  is a distinguishing feature of the family Tingidae. Due to their habitat, their diet consists of high quality sap which requires their mouth pieces to be both a sucker and a piercer.<ref name=":10" /> Tingidae have thin elongated legs with tarsi divided into two tarsomeres, allowing them to maneuver around the plant they are inhabiting.<ref name=":9" />

== Reproduction and development ==
Each individual usually completes its entire [[Biological life cycle|lifecycle]] on the same plant, if not the same part of the plant. The number of generations a species has annually is determined through photoperiodic induction.<ref name=":05" /> Most species have one to two generations per year, but some species have multiple generations. For example, all southern Indian Tingids are multivoltine, while most European species are univoltine.<ref name=":3">{{Cite journal |last1=Livingstone |first1=David |last2=Yacoob |first2=M H S |date=1987 |title=Biosystematics of Tingidae on the basis of the biology and micromorphology of their eggs |url=http://link.springer.com/10.1007/BF03179614 |journal=Proceedings: Animal Sciences |language=en |volume=96 |issue=5 |pages=587–611 |doi=10.1007/BF03179614 |issn=0253-4118|url-access=subscription }}</ref> The location where Tingidae species lay their eggs on the host plant varies, partially influenced by the specialization of their [[ovipositor]]. Eggs can be laid in the plant leaf lamina by being inserted horizontally to the leaf surface or by being inserted at an angle into the mesophyll. Oviposition also can occur in the leaf midrib, in floral parts such as the calyx or pistil, and in the stem.<ref name=":3" /> Tingidae lay dipause and non-cleidoic eggs.<ref name=":05" /> Due to the high water requirement for eggs to develop, the changes in moisture levels throughout seasons can impact the number of generations per year. A mechanism to cope with the fluctuations is to [[Overwintering|overwinter]] as eggs or nymphs; however, most overwinter as adults. Tingidae has incomplete [[metamorphosis]] in that the immature stages resemble the adults, except they are smaller and do not have wings. Maternal care of offspring is rare, however has been observed in some genera. For example, in ''Gargaphia solani,'' one female will brood not only its own eggs but also the eggs of other females, which as a result increases the survival rate of the offspring while reducing the number of females exposed to danger.<ref name=":05" />

== Habitat ==
[[File:Corythucha ciliata.jpg|thumb|253x253px|''Corythucha ciliata'' - Found by the river Neckar in Heidelberg, Germany.]]
Tingidae are generally [[Host (biology)|host]]-specific and are all completely phytophagous.<ref name=":05" /> Different species are dependent on a certain plant or a group of closely related plants for feeding. Different adaptations allows them to occupy a variety of different habitats. Habitats of multiple species can occur on a singular plant where different species can occupy anywhere between the roots and the top canopy.<ref name=":1">{{Cite journal |last1=Guilbert |first1=Eric |last2=Damgaard |first2=Jakob |last3=D'Haese |first3=Cyrille A. |date=2014 |title=Phylogeny of the lacebugs ( I nsecta: H eteroptera: T ingidae) using morphological and molecular data |url=https://resjournals.onlinelibrary.wiley.com/doi/10.1111/syen.12045 |journal=Systematic Entomology |language=en |volume=39 |issue=3 |pages=431–441 |doi=10.1111/syen.12045 |bibcode=2014SysEn..39..431G |issn=0307-6970|url-access=subscription }}</ref> With the majority of the members of family Tingidae having a sedentary lifestyle, many are found to inhabit the underside of leaves.<ref name=":05" /> Tingidae are able to withdraw sap from the plant by penetrating their stylet into the living plant tissue. Due to their feeding style, this method can be destructive to plants. The removal of sap from the plant cells causes the leaves to become discoloured and can eventually lead to wilted leaves.

Natural predators of Tingidae are not usually seen until populations become abundant. These predators include ladybugs, parasitic wasps, lacewings, and spiders.<ref name=":2">{{Cite journal |last1=Paulin |first1=Márton |last2=Hirka |first2=Anikó |last3=Eötvös |first3=Csaba Béla |last4=Gáspár |first4=Csaba |last5=Fürjes-Mikó |first5=Ágnes |last6=Csóka |first6=György |date=2020-11-01 |title=Known and predicted impacts of the invasive oak lace bug ( Corythucha arcuata ) in European oak ecosystems – a review |url=https://www.sciendo.com/article/10.2478/foecol-2020-0015 |journal=Folia Oecologica |language=en |volume=47 |issue=2 |pages=131–139 |doi=10.2478/foecol-2020-0015 |bibcode=2020FoOec..47..131P |issn=1338-7014|doi-access=free }}</ref> In Europe it was seen that natural predators increased when the population of ''Corythucha arcuata,'' oak lace bugs, increased. The natural predators of the oak lace bug located in Europe included lacewings, velvet mites and spiders. However even with the increase in predators, there was no significant decrease and regulation of the invasive oak lace bug.

For the species of Tingidae that inhabit the leaves of trees, some species of the genus ''Corythucha'' have been reported to land on humans and bite. Their bites may cause pain but are often not a concern to human health.<ref>{{Cite journal |date=2019-11-21 |title=Orius tristicolor (minute pirate bug) |url=https://doi.org/10.1079/cabicompendium.37841 |access-date=2025-03-24 |website=CABI Compendium|doi=10.1079/cabicompendium.37841 |url-access=subscription }}</ref> ''Corythucha ciliata'' bites have been reported in Europe in countries of Italy,<ref>{{Cite journal |last1=Maceljski |first1=Milan |last2=Balarin |first2=Inoslava |date=1977 |title=Beitrag zur Kenntnis natürlicher Feinde der Platanen-Netzwanze (Corythuca ciliata (Say), Tingidae, Heteroptera) |url=https://doi.org/10.1007/bf02156801 |journal=Anzeiger für Schädlingskunde Pflanzenschutz Umweltschutz |volume=50 |issue=9 |pages=135–138 |doi=10.1007/bf02156801 |bibcode=1977JPesS..50..135M |issn=0340-7330|url-access=subscription }}</ref> France<ref>{{Cite journal |last1=Izri |first1=Arezki |last2=Andriantsoanirina |first2=Valérie |last3=Chosidow |first3=Olivier |last4=Durand |first4=Rémy |date=2015-08-01 |title=Dermatosis Caused by Blood-Sucking Corythucha Ciliata |url=http://archderm.jamanetwork.com/article.aspx?doi=10.1001/jamadermatol.2015.0577 |journal=JAMA Dermatology |language=en |volume=151 |issue=8 |pages=909–910 |doi=10.1001/jamadermatol.2015.0577 |pmid=25970727 |issn=2168-6068|url-access=subscription }}</ref> and Romania.<ref>{{Cite journal |date=2021-11-16 |title=Corythucha (lace bugs) |url=https://doi.org/10.1079/cabicompendium.16262 |access-date=2025-03-24 |website=CABI Compendium|doi=10.1079/cabicompendium.16262 |url-access=subscription }}</ref> In some of these reported cases, the effects of ''C. ciliata'' bites have caused skin conditions such as dermatosis.

==Phylogeny==
Tingidae consist of roughly 2000 species in 3 subfamilies, Cantacadericies, Tingidae and Viananidinae.<ref name=":9" /> All 3 subfamilies can be found ubiquitous across the globe. The Cantacadericies are mainly concentrated all over the Southern Hemisphere, while Tinginae are located within tropical and temperate zones and Viananidinae in neotropical regions. Tinginae is the subfamily consistent of lace bugs, it includes 220 genera out of the 250 total identified Tingidae species. The higher order is as follows:

* Order: Hemiptera
** Suborder: Heteroptera
*** Superfamily: Tingoidea
**** Family: Tingidae
***** Subfamily: Cantacaderinae
***** Subfamily: Tinginae
***** Subfamily: Vianaidinae

Members have been found in the fossil record from the Early Cretaceous onwards, with the oldest being ''[[Sinaldocader]]'' from the Early Cretaceous [[Zaza Formation]] of [[Buryatia]], Russia.<ref>{{Cite journal |last1=Golub |first1=V. B. |last2=Popov |first2=Yu. A. |date=2008 |title=A new species of Tingidae (Insecta: Hemiptera: Heteroptera) from the Lower Cretaceous of Transbaikalia |url=https://link.springer.com/10.1134/S0031030108010140 |journal=Paleontological Journal |language=en |volume=42 |issue=1 |pages=86–89 |doi=10.1134/S0031030108010140 |bibcode=2008PalJ...42...86G |issn=0031-0301|url-access=subscription }}</ref><ref>{{Cite journal |last1=Guilbert |first1=Eric |last2=Heiss |first2=Ernst |date=2019 |title=New lacebugs (Hemiptera: Heteroptera: Tingidae) from Upper Cretaceous Burmese amber |url=https://linkinghub.elsevier.com/retrieve/pii/S0195667118301678 |journal=Cretaceous Research |language=en |volume=94 |pages=72–79 |doi=10.1016/j.cretres.2018.10.024|bibcode=2019CrRes..94...72G |url-access=subscription }}</ref> A new subfamily and genus of Tingidae was recently discovered in a piece of Burmese amber.<ref>{{Cite journal |last1=Heiss |first1=Ernst |last2=Golub |first2=Viktor B. |last3=Popov |first3=Yuri A. |date=2015 |title=A new subfamily, genus and species of Tingidae (Hemiptera: Heteroptera) from Burmese amber |url=https://www.zobodat.at/publikation_articles.php?id=257601 |journal=Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen |volume=67 |pages=1–9}}</ref> The preserved bug was assigned to the family Tingidae since it shared important morphological characteristics such as areolate hemelytra and paranota as well as 4-segmented antennae and labium. However, there were also significant differences that did not align with the previously established subfamilies such as the absence of spines on the head and a large scutellum. These differences helped lead to the recognition of the new subfamily Tingiometrinae and the species of the lace bug in the amber classified as ''Tingiometra burmanica.''

== Ecological importance ==
[[File:2021 07 27 Corythucha arcuata.jpg|thumb|302x302px|''Corythucha arcuata -'' Oak lace bug with eggs on the underside of an oak leaf]]
Lace bugs pose a potential threat to plants. Although not all species of lace bugs are pests, some species can affect various types of trees and crops of human consumption.<ref name=":05" /> ''Corythucha arcuata'' is a specific invasive Tingid that can cause significant damage to trees. ''C.arcuata'', native to North America, was first reported in Italy in 2002 and has since spread to multiple countries, causing considerable damage.<ref name=":4">{{Cite journal |last1=Bălăcenoiu |first1=Flavius |last2=Japelj |first2=Anže |last3=Bernardinelli |first3=Iris |last4=Castagneyrol |first4=Bastien |last5=Csóka |first5=György |last6=Glavendekić |first6=Milka |last7=Hoch |first7=Gernot |last8=Hrašovec |first8=Boris |last9=Krajter Ostoic |first9=Silvija |last10=Paulin |first10=Marton |last11=Williams |first11=David |last12=Witters |first12=Johan |last13=de Groot |first13=Maarten |date=2021 |title=Corythucha arcuata (Say, 1832) (Hemiptera, Tingidae) in its invasive range in Europe: perception, knowledge and willingness to act in foresters and citizens |url=https://neobiota.pensoft.net/article/71851/ |journal=NeoBiota |volume=69 |pages=133–153 |doi=10.3897/neobiota.69.71851 |doi-access=free |bibcode=2021NeoBi..69..133B |issn=1314-2488}}</ref> The pest causes chlorotic disease in oak along with the premature loss of leaves.<ref name=":5">{{Cite journal |last1=Kovač |first1=Marta |last2=Linde |first2=Andreas |last3=Lacković |first3=Nikola |last4=Bollmann |first4=Friedrich |last5=Pernek |first5=Milan |date=2021 |title=Natural infestation of entomopathogenic fungus Beauveria pseudobassiana on overwintering Corythucha arcuata (Say) (Hemiptera: Tingidae) and its efficacy under laboratory conditions |url=https://linkinghub.elsevier.com/retrieve/pii/S0378112721002814 |journal=Forest Ecology and Management |language=en |volume=491 |pages=119193 |doi=10.1016/j.foreco.2021.119193|bibcode=2021ForEM.49119193K |url-access=subscription }}</ref> Furthermore, there have been reports of bites and skin irritations from the species in urban areas. Insecticides have been used to keep ''C.arcruata'' at bay, however, this can result in unfavourable effects on the surrounding environment. Mechanical control methods, including the use of yellow sticky traps and suction traps, have also been employed.<ref name=":4" />

Tingidae can act as possible biological control agents when controlling specific weed populations. In New Zealand, the invasive weed ''Solanum mauritianum'', also known as woolly nightshade can be controlled by releasing ''Gargaphia decoris''.<ref name=":6">{{Cite journal |last1=Falla |first1=Cecilia |last2=Minor |first2=Maria |last3=Harrington |first3=Kerry |last4=Paynter |first4=Quentin |last5=Cordiner |first5=Sarah |last6=Najar-Rodriguez |first6=Adriana |date=2023-06-01 |title=Effects of light intensity on Solanum mauritianum (Solanaceae) morphological and chemical traits and the performance of its biological control agent Gargaphia decoris (Hemiptera: Tingidae) |url=https://linkinghub.elsevier.com/retrieve/pii/S1049964423000713 |journal=Biological Control |volume=181 |pages=105218 |doi=10.1016/j.biocontrol.2023.105218 |bibcode=2023BiolC.18105218F |issn=1049-9644|url-access=subscription }}</ref> Higher numbers of ''G. decoris'' were spotted on the leaves of woolly nightshade when grown in the shade when compared to ones growing in full sunlight. ''G. decoris'' causes damage by removing chlorophyll from the leaves, which ultimately decreases the rate of photosynthesis of the woolly nightshade.<ref name=":7">{{Cite journal |last1=Cowie |first1=Blair W. |last2=Byrne |first2=Marcus J. |last3=Witkowski |first3=Ed T. F. |last4=Venter |first4=Nic |date=2016-03-01 |title=Exacerbation of photosynthetic damage through increased heat–light stress resulting from Gargaphia decoris sap-feeding |url=https://linkinghub.elsevier.com/retrieve/pii/S1049964415300669 |journal=Biological Control |volume=94 |pages=82–89 |doi=10.1016/j.biocontrol.2015.12.008 |bibcode=2016BiolC..94...82C |issn=1049-9644|url-access=subscription }}</ref> ''Gargaphia decoris'' sap-feeding method can further reduce photosynthesis by reducing the diffusion of gases through the plants stomata. In addition, the reduced rates of stomatal conductance inhibits ''Solanum mauritianum'' ability to participate in evaporative cooling leading to a greater damaging effect, especially in the leaves that received full sunlight. The impact of ''G. decoris'' on controlling the invasive species is greatest through the combination of their feeding methods and indirectly through factors, like accumulated heat from sunlight, that cause physiological impairment.
==Gallery==
{{gallery|File:Physatocheila smreczynskii (Tingidae), Nijkerk, the Netherlands.jpg|''Physatocheila smreczynskii''|File:Cochlochila species lace bug South Africa composite.jpg|Adult specimen of a small (about 2 mm) species of lace bug on ''[[Lavender|Lavandula]]'' near Cape Town in South Africa: Left, dorsal view; right, lateral view, showing proboscis and dorsal protuberances|File:Gargaphia P1220765a.jpg|''[[Gargaphia]]'' from [[Oklahoma]]|File:Tingidae.jpg|from [[Maryland]]|File:Lace bug - Pennsylvania.jpg|alt=|Lace bug - Pennsylvania|width=250|File: Tingidae - Lace Bug.png|Lace bug|Dictyonota fuliginosa (Tingidae) - (imago), Molenhoek, the Netherlands - 3.jpg|Dictyonota fuliginosa}}
==See also==
*''[[Gargaphia solani]]'' – eggplant lacebug
*''[[Stephanitis takeyai]]'' – andromeda lace bug
*{{extinct}}''[[Leptopharsa tacanae]]'' - tingid from [[Mexican amber]]

==References==
{{Reflist}}

==Further reading==
{{refbegin}}
*Ciceoi, R. & Radulovici, A. Faculative blood-sucking lace bugs, Corythucha sp., in Romania.
*Drake, C.J. & Ruhoff, F.A., 1960. Lace-bug genera of the world. (Hemiptera: Tingidae). Proc. U.S. Natl. Mus. 112 (3431): 1–105, 9 pls.
*DUTTO, M.& BERTERO, M. 2013. Dermatosis caused by Corythuca ciliata (Say, 1932) (Heteroptera, Tingidae). Diagnostic and clinical aspects of an unrecognized pseudoparasitosis. J Prev Med Hyg. 54 (1): 57-59
*Froeschner, R.C., 1996. Lace Bug Genera of the World, I: Introduction, Subfamily Canthacaderinae (Heteroptera: Tingidae). Smithsonian Contributions to Zoology, No. 574.
*Froeschner, R.C., 2001. Lace Bug Genera of the World, II: Subfamily Tinginae: tribes Litadeini and Ypsotingini (Heteroptera: Tingidae). Smithsonian Contributions to Zoology, No. 611. 
*Javahery, J. 2019. Natural history of Corythucha juglandis and Gargaphia tiliae (Hemiptera: Tingidae) in Ontario and Québec, Canada. The Canadian Entomologist. 151 (4): 475–497.
*Lis, Barbara; Zielińska, Anna; Lis, Jerzy A. 2022. The King's Lace Bug Recaredus rex Distant, 1909 (Hemiptera: Heteroptera: Tingidae): Systematic Position, First Palaearctic and Afrotropical Records, and Ecological Niche Modelling". Insects. 13 (6): 558.{{Refend}}

==External links==
{{Wikispecies|Tingidae}}
{{Commons category|Tingidae}}
*[http://www.britishbugs.org.uk/gallery/heteroptera/Tingidae/tingidae.html Lacebugs of Britain]
*[http://entomology.ifas.ufl.edu/fasulo/woodybug/lacebugs.htm lace bugs of southeastern U.S. woody ornamentals]
*On the [[University of Florida]] / [[Institute of Food and Agricultural Sciences]] ''Featured Creatures'' website
:*[http://entomology.ifas.ufl.edu/creatures/trees/sycamore_lace_bug.htm ''Corythucha ciliata'', sycamore lace bug]
:*[http://entomology.ifas.ufl.edu/creatures/orn/hawthorn_lace_bug.htm ''Corythucha cydoniae'', hawthorn lace bug] 
:*[http://entomology.ifas.ufl.edu/creatures/field/sugarcane_lace_bug.htm ''Leptodictya tabida'', sugarcane lace bug] 
:*[http://entomology.ifas.ufl.edu/creatures/fruit/avocado_lace_bug.htm ''Pseudacysta perseae'', avocado lace bug] 
:*[http://entomology.ifas.ufl.edu/creatures/orn/shrubs/azalea_lace_bug.htm ''Stephanitis pyrioides'', azalea lace bug] 
:*[http://entomology.ifas.ufl.edu/creatures/beneficial/lantana_lace_bug.htm ''Teleonemia scrupulosa'', lantana lace bug]

{{Hemiptera|3}}

{{Taxonbar|from=Q1472961}}
{{Authority control}}

[[Category:Tingidae| ]]
[[Category:Heteroptera families]]