ZooKeys 319: 169–189, doi: 10.3897/zookeys.319.4316
True bugs (Hemiptera, Heteroptera) as psyllid predators (Hemiptera, Psylloidea)
Dušanka Jerinić-Prodanović 1, Ljiljana Protić 2
1 University of Belgrade, Faculty of Agriculture, Nemanjina 6, 11080 Zemun, Serbia
2 Natural History Museum, Njegoševa 51, 11000 Belgrade, Serbia

Corresponding author: Dušanka Jerinić-Prodanović (dusanka@agrif.bg.ac.rs)

Academic editor: A. Popov

received 14 November 2012 | accepted 28 May 2013 | Published 30 July 2013


(C) Dušanka Jerinić-Prodanović. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


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Citation: Jerinić-Prodanović D, Protić L (2013) True bugs (Hemiptera, Heteroptera) as psyllid predators (Hemiptera, Psylloidea). In: Popov A, Grozeva S, Simov N, Tasheva E (Eds) Advances in Hemipterology. ZooKeys 319: 169–189. doi: 10.3897/zookeys.319.4316

Abstract

Data on natural enemies of psyllids are rare and can usually be found in papers about economically significant species. During an investigation of psyllid fauna in Serbia, natural enemies were investigated, too. True bugs were the most numerous among them. From 28 psyllid species, 21 species of true bugs from families Anthocoridae and Miridae were reared. Seven species of Anthocoridae were identified: Anthocoris amplicollis (Horváth, 1839), Anthocoris confusus Reuter, 1884, Anthocoris nemoralis (Fabricius, 1794), Anthocoris nemorum (Linnaeus, 1761), Orius majusculus Reuter, 1884, Orius minutus (Linnaeus, 1758) and Orius niger Wolff, 1811. The following 14 species of Miridae were identified: Atractotomus mali Meyer-Dür, 1843, Campylomma verbasci (Meyer-Dür, 1843), Deraeocoris flavilinea (A. Costa, 1862), Deraeocoris ruber (Linnaeus, 1758), Deraeocoris lutescens (Schilling, 1836), Heterocordylus genistae (Scopoli, 1763), Hypseloecus visci (Puton, 1888), Malacocoris chlorizans Panzer, 1794, Miris striatus (Linnaeus, 1758), Orthotylus marginalis Reuter, 1884, Psallus assimilis Stichel, 1956, Psallus quercus Kirschbaum, 1856, Psallus flavellus Stichel, 1933 and Pseudoloxops coccinea (Meyer-Dür, 1843). The aim of the research was to provide list of true bugs recorded as predators of psyllids in order to preserve their diversity and significance, especially on cultivated plants.

Keywords

Psylloidea, Heteroptera, predators, natural enemies, Serbia

Introduction

Predators of psyllids (Psylloidea) have been poorly known. So far, detailed researches were carried out only on the predators of economically significant species, such as pear psyllids Cacopsylla pyri (Linnaeus, 1758), Cacopsylla pyricola (Foerster, 1848) and Cacopsylla pyrisuga (Foerster, 1848); apple psyllid Cacopsylla mali (Schmidberger, 1836) and eucalyptus psyllids from the subfamily Spondyliaspidinae (Jonsson 1983, Herard 1985, 1986, Santas 1987, Erler 2004, Horton et al. 2004, Sigsgaard et al. 2006, Jauset et al. 2006, Luiz de Queiroz et al. 2012). There are too few data on predators of other psyllid species. Hodkinson and Flint (1971) investigated predators of ash psyllid, Psyllopsis fraxini (Linnaeus, 1758), in England, and Harizanova et al. (2012), predatory complex of Acizzia jamatonica (Kuwayama, 1908) in Bulgaria. In these papers, the most represented are psyllid predators from the order Hemiptera (suborder Heteroptera) followed by Coleoptera, Neuroptera, Diptera, Dermaptera and Acari. Within the Heteroptera, the most numerous in species families are Anthocoridae, Miridae and Nabidae. A polyphagous species, Anthocoris nemoralis (Fabricius, 1794), was most frequently found, with a preference for the species from superfamily Psylloidea (Jonsson 1983, Herard 1986). Anthocoris nemoralis (Anthocoridae) was introduced from Europe to North America (British Columbia) in 1963 in order to control Cacopsylla pyricola, where its establishment was successful. Besides giving satisfactory effects, this species also spread in the new environment suppressing autochthonous species Anthocoris antevolens White, 1879 and Anthocoris melanocerus Reuter, 1884, which are most common anthocorid predators in orchards (Herard 1986, Horton et al. 2004).

Data on psyllid predators in Serbia relate only to the predators of pear psyllids (Pavićević 1977, Grbić et al. 1989, Jerinić-Prodanović et al. 2010).

Pavićević (1977) found a large number of predatory species, among which two were from family Anthocoridae. Grbić et al. (1989) recorded four species of Heteroptera: Anthocoris nemoralis and Orius sp. (both Anthocoridae), Pilophorus clavatus (Linnaeus, 1767) (Miridae) and Nabis pseudoferus Remane, 1949 (Nabidae), while Jerinić-Prodanović et al. (2010) reported seven species: Anthocoris nemoralis (Fabricius, 1794), Anthocoris nemorum (Linnaeus, 1761), Orius (Heterorius) minutus (Linnaeus, 1758) and Orius (Orius) niger Wolff, 1811 from the family Anthocoridae and Campylomma verbasci (Meyer-Dür, 1843), Deraeocoris (Deraeocoris) ruber (Linnaeus, 1758) and Deraeocoris (Knightocapsus) lutescens (Schilling, 1836) from the family Miridae.

There is no data on other predatory psyllid species in Serbia.

Methods

Insect material was collected from 419 localities within the whole territory of the Republic of Serbia. Investigations were carried out in the period from 2005 to 2010, in field conditions and in the laboratory of the Faculty of Agriculture in Zemun, University of Belgrade. Locality mapping was carried out in World UTM (Universal Transverse Mercator) cartographic projection. Determination of coordinates of investigated localities in the field was carried out using GPS devices Geoexplorer 3 (Trimble) and E-trex Vista Hcx (Garmin), with an accuracy of 3 to 5 meters.

Adults of predatory true bugs were collected from psyllid colonies by an aspirator and their larvae were collected together with plant material and psyllids and further reared to adults in laboratory conditions in Petri dishes.

The species identification of Heteroptera was based on Wagner (1970–1971, 1975), Péricart (1972) and Kerzhner and Josifov (1999).

A part of the material is deposited in the first author’s collection in the Faculty of Agriculture, University of Belgrade, and another part, in the second author’s collection in Natural History Museum, Belgrade.

Results and discussion

We collected and reared 21 true bug species predating on 28 psyllid species (Table 1) from 44 localities (Fig. 1 and Table 2). The identified true bugs belong to families Anthocoridae and Miridae.

Table 1.

List of preys (Psylloidea) and their predators (Heteroptera) and host plants in Serbia.

Preys (Psylloidea) Predators (Heteroptera)
Anthocoridae Miridae
Ant amp Ant con Ant nea Ant neu Ori maj Ori min Ori nig Atr mal Cam ver Der fla Der lut Der rub Het gen Hyp vis Mal chl Mir str Ort mar Psa ass Psa fla Psa que Pse coc
Baeopelma foersteri Aln glu
Cacopsylla affinis Cra mon Cra mon
Cacopsylla bidens Pyr com Pyr com Pyr com Pyr com Pyr com
Cacopsylla melanoneura Cra mon Mal dom Mal dom Mal dom,
Cra mon
Mal dom Mal dom,
Cra mon
Cra mon
Cacopsylla peregrina Cra mon Cra mon Cra mon
Cacopsylla picta Mal dom Mal dom
Cacopsylla pulchra Sal pur
Cacopsylla pyri Pyr com Pyr com Pyr com Pyr com Pyr com Pyr com Pyr com
Cacopsylla pyricola Pyr com
Cacopsylla pyrisuga Pyr com Pyr com Pyr com
Cacopsylla rhamnicola Rha cat Rha cat Rha cat
Cacopsylla visci Vis alb Vis alb Vis alb
Camarotoscena speciosa Pop nig Pop nig Pop nig
Craspedolepta sp. Art vul
Homotoma ficus Fic car Fic car Fic car
Livia junci Jun bul
Psylla buxi Bux sem Bux sem Bux sem
Psyllopsis discrepans Fra spp Fra spp Fra orn Fra ang Fra spp Fra spp Fra orn,
Fra spp
Psyllopsis fraxini Fra spp
Psyllopsis fraxinicola Fra spp Fra ang Fra spp Fra orn Fra ang Fra spp Fra spp
Psyllopsis machinosa Fra spp Fra spp Fra orn Fra ang Fra spp
Psyllopsis meliphila Fra spp
Psyllopsis repens Fra spp Fra orn
Trichochermes walkeri Rha cat
Trioza chenopodii Atr tat Atr obl
Trioza mesembrina Cha hir
Trioza rhamni Rha cat Rha cat
Trioza urticae Urt dio Urt dio

Abbreviations
Predators
. Anthocoridae: Ant amp, Anthocoris amplicollis; Ant con, Anthocoris confusus; Ant nea, Anthocoris nemoralis; Ant neu, Anthocoris nemorum; Ori maj, Orius majusculus; Ori min, Orius minutus; Ori nig, Orius niger; Miridae: Atr mal, Atractotomus mali; Cam ver, Campylomma verbasci; Der fla, Deraeocoris flavilinea; Der lut, Deraeocoris lutescens; Der rub, Deraeocoris ruber; Het gen, Heterocordylus genistae; Hyp vis, Hypseloecus visci; Mal chl, Malacocoris chlorizans; Mir str, Miris striatus; Ort mar, Orthotylus marginalis; Psa ass, Psallus assimilis; Psa fla, Psallus flavellus; Psa que, Psallus quercus; Pse coc, Pseudoloxops coccinea.
Host plants: Aln glu, Alnus glutinosa; Art vul, Artemisia vulgaris; Atr obl, Atriplex oblongifolia; Atr tat, Atriplex tatarica; Bux sem, Buxus sempervirens; Cha hir, Chaerophyllum hirsutum; Cra mon, Crataegus monogyna; Fic car, Ficus carica; Fra ang, Fraxinus angustifolia; Fra orn, Fraxinus ornus; Fra spp, Fraxinus spp.; Jun bul, Juncus bulbosus; Mal dom, Malus domestica; Pop nig, Populus nigra; Pyr com, Pyrus communis; Rha cat, Rhamnus cathartica; Sal pur, Salix purpurea; Vis alb, Viscum album; Urt dio, Urtica dioica.

Table 2.

Geographical coordinates of inspected localities.

No Locality Latitude, Longitude Altitude
1 Bački breg 45°55'21"N, 18°55'24"E 90
2 Bavanište 44°48'42"N, 20°53'10"E 80
3 Beloljin 43°14'03"N, 21°24'26"E 290
4 Beograd–Autokomanda 44°47'20"N, 20°28'20"E 100
5 Beograd–Banjica 44°45'18"N, 20°28'58"E 190
6 Beograd–Block 45 44°47'36"N, 20°22'47"E 75
7 Beograd–Bulevar Aleksandra Karadjordjevića 44°46'50"N, 20°27'31"E 175
8 Beograd–Hotel Jugoslavija 44°49'36"N, 20°25'22"E 75
9 Beograd–Hram Svetog Save 44°47'53"N, 20°28'03"E 120
10 Beograd–Kalemegdan 44°49'19"N, 20°26'52"E 110
11 Beograd–Karaburma 44°48'48"N, 20°29'15"E 110
12 Beograd–Milošev konak 44°46'38"N, 20°25'36"E 80
13 Beograd–Voždovac 44°47'17"N, 20°28'28"E 85
14 Brestovačka Banja 44°03'36"N, 22°02'36"E 360
15 Dobra 44°38'23"N, 21°54'06"E 85
16 Draževac 43°28'08"N, 21°46'37"E 205
17 Galovica 44°46'22"N, 20°21'04"E 75
18 Grocka 44°41'21"N, 20°42'02"E 125
19 Ilinci 45°06'41"N, 19°07'16"E 80
20 Izvor 43°04'58"N, 22°23'57"E 290
21 Kelebija 46°08'59"N, 19°35'10"E 125
22 Kopaonik–Srebrenac 43°19'02"N, 20°50'08"E 1740
23 Klokočevac 44°20'53"N, 22°10'45"E 140
24 Koruška 45°11'46"N, 19°34'23"E 110
25 Lipovača 45°08'24"N, 19°16'53"E 165
26 Luka 44°09'46"N, 22°11'56"E 340
27 Majdanpek 44°25'10"N, 21°57'10"E 520
28 Nemenikuće 44°29'38"N, 20°34'00"E 280
29 Niš 43°18'35"N, 21°53'50"E 200
30 Novi Sad–Detelinara 45°15'50"N, 19°48'56"E 80
31 Obedska bara 44°44'10"N, 19°59'15"E 80
32 Oparić 43°44'40"N, 21°06'38"E 310
33 Radenković 44°56'01"N, 19°29'05"E 80
34 Radmilovac 44°45'15"N, 20°34'39"E 160
35 Sokobanja 43°38'41"N, 21°53'11"E 350
36 Sutjeska 45°23'02"N, 20°41'53"E 75
37 Šid 45°07'31"N, 19°12'58"E 105
38 Umčari 44°34'10"N, 20°43'00"E 160
39 Uzovnica 44°16'12"N, 19°19'47"E 170
40 Veliko Središte 45°12'54"N, 21°25'30"E 120
41 Vrujci 44°13'26"N, 20°09'53"E 170
42 Zemun–Nova Galenika 44°51'41"N, 20°22'11"E 90
43 Zemunski kej 44°50'29"N, 20°25'06"E 75
44 Zlatibor–Kraljevske Vode 43°43'39"N, 19°42'06"E 950
Figure 1.

Localities in Serbia where true bug predators of psyllids were collected.

Anthocoridae
1) Anthocoris amplicollis (Horváth, 1839)
Trophic status.

Zoophagous.

Distribution.

Europe.

Prey.

Psylla buxi (Linnaeus, 1758), from Buxus sempervirens, Nova Galenika, 13.VI.2009, reared 3♂♂, 2♀♀.

Anthocoris amplicollis was already reported in Serbia (Protić and Stojanović 2003) but the above mentioned record is the first one in Serbia as a psyllid predator. In Switzerland, it was registered by Wyniger and Burckhardt (2003) in galls of Psyllopsis fraxini. According to available literature data, Anthocoris amplicollis has not been published as a predator of Psylla buxi.

2) Anthocoris confusus Reuter, 1884
Trophic status.

Zoophagous.

Distribution.

Palaearctic.

Prey.

Psylla buxi, from Buxus semprevirens, Sokobanja, 25.IX.2009, reared 1♀.

Registered as a psyllid predator on conifers (Wyniger and Burckhardt 2003) and aphids (Herard 1986). In the present paper, reported for the first time as a psyllid predator in Serbia.

3) Anthocoris nemoralis (Fabricius, 1794)
Trophic status.

Zoophagous.

Distribution.

Euro-Mediterranean.

Preys.

Cacopsylla bidens (Šulc, 1907), from Pyrus communis, Beograd–Karaburma, 19.V.2006, reared 2♀♀ ‘ex larva' 25.V.2006. Cacopsylla pyri, from Pyrus communis, Nemenikuće, 15.VI.2006, reared 1♂ ‘ex larva' 22.VI.2006; Radmilovac, 2.IX.2005, reared 1♂; 12.VI.2006, reared 1♂ ‘ex larva' 25.VI.2006; 26.X.2006, collected 1♂; 7.VI.2007, reared 1♂, 1♀ ‘ex larva' 11.VI.2007, 20.VI.2007, collected 1 larva and 1♂; 26.X.2008, reared 1♂ ‘ex larva' 3.XI.2008. Cacopsylla pyrisuga, from Pyrus communis, Zemunski kej, 28.V.2007, collected 1♀. Cacopsylla visci (Curtis, 1835), from Viscum album, Beograd–Bulevar Aleksandra Karađorđevića, 25.III.2007, reared 2♂♂, 1♀ ‘ex larva' 12.IV.2007; 23.IV.2007, reared 2♂♂, 5♀♀ ‘ex larva' 3.V.2007. Camarotoscena speciosa (Flor, 1861), from Populus nigra, Radmilovac, 22.V.2000, reared 1♂ ‘ex larva' 25.V.2000; Zemun–Nova Galenika, 25.VIII.2008, collected 1♀; 1.IX.2008, reared 3♀♀ ‘ex larva' 9.IX.2008; 10.IX.2008, collected 1♀, 1♂; 17.IX.2008, collected 1♀, 1♂; 24.IX.2008, reared 2♂♂, 2♀♀ ‘ex larva' 1.X.2008; 27.VII.2010, collected 1♂, 1♀, 18.X.2010, collected 1♂, 1♀. Psyllopsis discrepans (Flor, 1861), from Fraxinus sp., Beograd–Autokomanda, 10.V.2007, reared 1♀; Brestovačka Banja, 25.V.2007, reared 5♂♂, 4♀♀ ’ex larva'8.VI.2007; Ilinci, 24.V.2008, collected 1♂; Majdanpek, 25.V.2007, reared 1♂, 1♀ ‘ex larva' 3.VI.2007, Milošev konak, 21.VI.2007, 1♂. Psyllopsis fraxini, from Fraxinus sp., Veliko Središte, 30.V.2006, reared 2♀♀. Psyllopsis fraxinicola (Foerster, 1848), from Fraxinus sp., Beograd–Autokomanda, 10.V.2007, reared 1♀; Beograd–Kalemegdan, 24.V.2007, collected 1♂; Brestovačka Banja, 25.V.2007, reared 5♂♂, 4♀♀ ‘ex larva' 8.VI.2007; Majdanpek, 25.V.2007, reared 1♂, 1♀ ‘ex larva' 3.VI.2007; Veliko Središte, 4.VI.2006, reared 2♀♀ ‘ex larva' 19.VI.2006. Psyllopsis machinosa Loginova, 1963, from Fraxinus sp., Beograd–Autokomanda, 10.V. 2007, reared 1♀. Psyllopsis meliphila Löw, 1881, from Fraxinus sp., Nemenikuće, 15.VI.2006, collected 1♂. Psyllopsis repens Loginova, 1963, from Fraxinus sp., Beograd–Autokomanda, 14.X.2008, reared 1♀. Trioza urticae (Linnaeus, 1758), from Urtica dioica, Ilinci, 24.VI.2007, collected 2♀♀.

Anthocoris nemoralis is an important component of the natural enemy community in pear and apple orchards where it provides biological control against arthropod pests, particularly psyllids (Horton et al. 2004). Investigating the predator–prey complex of Cacopsylla pyrisuga in a pear orchard in France, Herard (1986) found that Anthocoris nemoralis was the most efficient enemy against this pest. Anthocoris nemoralis is mentioned in many papers as a permanent member of biocomplexes of pear psyllids in Europe (Wheeler 2000b, Erler 2004, Sigsgaard et al. 2006). In Turkey, Anthocoris nemoralis was an equally present and efficient predator of pear psyllid Cacopsylla pyrisuga, both in treated and untreated orchards, but still insufficient for its full control (Erler 2004). In Spain, Jauset et al. (2006) determined Anthocoris nemoralis as a very efficient predator of Cacopsylla pyrisuga, both in treated and untreated pear orchards. Now, there is a mass production of Anthocoris nemoralis in companies specialized for biological control of harmful insects (Sigsgaard et al. 2006). The same authors reported that Anthocoris nemoralis mostly prefers Cacopsylla pyrisuga to aphids, and that it prefers laying eggs on pear to apple. Anthocoris nemoralis is a polyphagous predatory species having psyllids as a usual prey.

It is distributed in Europe and the Mediterranean. From Europe it was introduced into North America in 1963 in order to control pear psyllid Cacopsylla pyricola, giving satisfactory results (Horton et al. 2004). This species has adapted to this region so well that it has suppressed autochthonous predatory species Anthocoris antevolens and Anthocoris melanocerus (Herard 1986, Horton et al. 2004). In Serbia, in a pear orchard, Pavićević (1977), Grbić et al. (1989) and Jerinić-Prodanović (2010) note a permanent presence of Anthocoris nemoralis, both during vegetation and winter period together with an overwintering adult of Cacopsylla pyrisuga. Anthocoris nemoralis was also reported as a predator of Psyllopsis repens in Serbia (Malenovský and Jerinić-Prodanović 2011).

4) Anthocoris nemorum (Linnaeus, 1761)
Trophic status.

Zoophagous.

Distribution.

Eurosiberian.

Preys.

Cacopsylla affinis (Löw, 1880), from Crataegus monogyna, Ilinci, 27.IV.2008, reared 1♂ ‘ex larva' 14.V. 2008. Cacopsylla bidens, from Pyrus communis, Nemenikuće, 15.VI.2006, reared 1♀. Cacopsylla melanoneura (Foerster, 1848), from Crataegus monogyna, Ilinci, 27.IV.2008, reared 1♂ ‘ex larva' 14.V.2008; Klokočevac, 10.V.2008, collected 1♀. Cacopsylla peregrina (Foerster, 1848), from Crataegus monogyna, Ilinci, 27.IV.2008, reared 1♂ ‘ex larva' 14.V.2008. Cacopsylla pulchra (Zetterstedt, 1838), from Salix purpurea, Zlatibor–Kraljevske Vode, 30.IV.2007, collected 1♀ and 1 larva. Cacopsylla pyri, from Pyrus communis, Novi Sad–Detelinara, 23.V.2008, reared 2♂♂, 3♀♀ ‘ex larva' 4.VI.2008; 14.VI.2008, reared 3♂♂ ‘ex larva' 17.VI.2008. Cacopsylla pyrisuga, from Pyrus communis, Grocka, 10.V.2008, reared 1♀ ‘ex larva' 23.V.2008. Cacopsylla visci, from Viscum album, Beograd–Bulevar Aleksandra Karađorđevića, 23.IV.2007, reared 1♂ ‘ex larva' 3.V.2007. Psylla buxi, from Buxus sempervirens, Vrujci, 1.VI.2009, collected 2♂♂, 1♀; Šid, 3.V.2008, collected 2♀♀; Zemun–Nova Galenika, 4.V.2008, reared 2♂♂, 3♀♀ ‘ex larva' 15.V.2008; 30.VII.2008, collected 1♀; 14.VI.2009, collected 2 specimens. Psyllopsis fraxinicola, from Fraxinus angustifolia, Beograd–Hram Svetog Save, 14.IV.2008, reared 1♂ and 1 larva.

Anthocoris nemorum is noted as a predator of many insect species, in the first place Hemiptera, Diptera, eggs of Lepidoptera and mites (Herard 1986, Wheeler 2000b, Sigsgaard et al. 2006), already registered as a predator of both Cacopsylla pyrisuga and Psyllopsis fraxini (Herard 1986). It is also largely reported as an efficient predator of apple psyllid Cacopsylla mali in Norway (Jonsson 1983). In England, Hodkinson and Flint (1971) determined Anthocoris nemorum as a predator of Psyllopsis fraxini collected from ash, while in Germany Novak and Achtziger (1995) registered it as a predator of hawthorn psyllids Cacopsylla melanoneura and Cacopsylla peregrina. Sigsgaard et al. (2006) note Anthocoris nemorum as a more polyphagous species than Anthocoris nemoralis. They also determined in experimental conditions that Anthocoris nemorum prefers aphids to psyllids, and has a preference for laying eggs on apple rather than on pear.

Anthocoris nemorum is an Eurosiberian species, introduced to North America in order to control Cacopsylla pyricola just like Anthocoris nemoralis, but without satisfactory results (Herard 1986).

Anthocoris nemorum is reported here for the first time as a predator of psyllids in Serbia.

5) Orius (Heterorius) majusculus Reuter, 1884
Trophic status.

Zoophagous.

Distribution.

Euro-Atlantic.

Preys.

Psyllopsis discrepans and Psyllopsis machinosa, from Fraxinus spp., Beograd–Autokomanda, 6.V.2009, collected 1♂.

Orius majusculus was registered as a predator of psyllids (Herard 1986). It is noted as a predator of aphids, such as Diuraphis noxia and Schizaphis graminum in Russia, mites in France, whiteflies in greenhouses in Italy and pear psyllid Cacopsylla pyri in France (Péricart 1972, Herard 1986).

The present paper reports Orius majusculus as a psyllid predator for the first time in Serbia and Psyllopsis discrepans and Psyllopsis machinosa for the first time as a prey of Orius majusculus.

6) Orius (Heterorius) minutus (Linnaeus, 1758) Fig. 2
Trophic status.

Zoophagous.

Distribution.

Palaearctic.

Preys.

Baeopelma foersteri (Flor, 1861), from Alnus glutinosa, Radenković, 3.VI.2006, collected 1♀. Cacopsylla melanoneura, from Malus domestica, Beograd–Hotel Jugoslavija, 26.V.2005, reared 1♂; Ilinci, 2.V.2010, reared 1♂, 2♀♀ ‘ex larva' 10.V.2010. Cacopsylla picta (Foerster, 1848), from Malus domestica, Beograd–Hotel Jugoslavija, 26.V.2005, reared 1♂. Cacopsylla pyri, from Pyrus communis, Radmilovac, 10.VII.2006, reared 1♂, 2♀♀ ‘ex larva' 20.VII.2006; 26.VII.2006, collected 1♂; 4.IX.2006, collected 1♂. Cacopsylla rhamnicola (Scott, 1876), from Rhamnus cathartica, Kelebija, 25.V.2005, reared 1♀ ‘ex larva' 6.VI.2005. Camarotoscena speciosa, from Populus nigra, Zemun–Nova Galenika, 1.IX.2008, reared 1♂ ‘ex larva' 9.IX.2008; 24.IX.2008, reared 1♂ ‘ex larva' 1.X.2008. Homotoma ficus (Linnaeus, 1758), from Ficus carica, Beograd–Banjica, 23.IX.2008, collected 2♂♂, 3♀♀, feeding on eggs. Psyllopsis discrepans, from Fraxinus ornus, Ilinci, 21.V.2005, collected 1♂. Psyllopsis discrepans and Psyllopsis repens, from Fraxinus ornus, Beograd–Autokomanda, 7.IX.2008, reared 5♂♂, 4♀♀ ‘ex larva' 11.IX.2008; 21.IX.2008, collected 3♂♂; 23.IX.2008, collected 1♂; 7.X.2008, collected 1♂, 1♀; 14.X.2008, collected 1♀; 21.X.2008, collected 1♀. Psyllopsis fraxinicola, from Fraxinus sp., Vrujci, 1.VII.2009, 1♀. Psyllopsis machinosa, from Fraxinus ornus, Beograd–Autokomanda, 6.V.2009, reared 1♀ ‘ex larva' 10.V.2009. Psyllopsis repens, from Fraxinus ornus, Beograd–Autokomanda, 21.VIII.2010, reared 1♀. Trioza mesembrina Burckhardt, 1986, from Chaerophyllum hirsutum, Kopaonik–Srebrenac, 7.VIII.2008, reared 1♂. Trioza rhamni (Schrank, 1801), from Rhamnus cathartica, Kelebija, 25.V.2005, reared 1♀ ‘ex larva' 6.VI.2005; Ilinci, 2.V.2009, 1♂.

Orius minutus is an extremely polyphagous species distributed in Europe, Siberia, China and Mediterranean region. Many authors determined it as a predator of harmful insect species from a number of orders (Thysanoptera, Diptera, Lepidoptera, Coleoptera and Hemiptera Homoptera). Already reported as a psyllid predator (Herard 1986). In France, Herard (1986) determined Orius minutus as a predator of pear psyllids, primarily Cacopsylla pyrisuga, and hawthorn psyllids. Also in Slovenia, Vrabl and Matis (1977) register it as a predator of Cacopsylla pyrisuga and Cacopsylla pyrisuga. In Serbia, Pavićević (1977) and Jerinić-Prodanović et al. (2010) determined Orius minutus as a predator of Cacopsylla pyrisuga in pear orchards. Malenovský and Jerinić-Prodanović (2011) also found it as a predator of Psyllopsis repens. In Croatia, Arčanin and Balarin (1972) recognized the significance of Orius minutus in the reduction of the mite Panonychus ulmi.

Figure 2.

Orius minutus feeding on Trioza rhamni.

7) Orius (Orius) niger Wolff, 1811
Trophic status.

Zoophagous.

Distribution.

Palaearctic.

Preys.

Cacopsylla bidens, from Pyrus communis, Ilinci, 13.X.2008, collected 1♂. Cacopsylla melanoneura, from Malus domestica, Lipovača, 29.IV.2006, collected 1♂. Cacopsylla pyri, from Pyrus communis, Radmilovac, 26.VI.2006, reared 1♂ ‘ex larva' 30.VI.2006; 10.VII.2006, reared 1♀ ‘ex larva' 20.VII.2006. Craspedolepta sp., from Artemisia vulgaris, Sutjeska, 2.X.2009, reared 1♂ and 4♀♀. Psyllopsis fraxinicola, from Fraxinus ornus, Vrujci, 30.VI.2009, collected 1 specimen. Livia junci (Schrank, 1789), from Juncus bulbosus, Beograd–Block 45, 10.VIII.2005, reared 1♂ ‘ex larva' 16.VIII.2005. Trichochermes walkeri (Foerster, 1848), from Rhamnus cathartica, Ilinci, 14.IX.2008, 2♂♂, 2♀♀. Trioza chenopodii Reuter, 1876, from Atriplex tatarica, Ilinci, 20.VIII.2006, 1♂ ‘ex larva' 23.VIII.2006. Trioza urticae, from Urtica dioica, Bački breg, 7.VI.2005, reared 1♂; Ilinci, 14.V.2005, reared 1♂, 1♀ ‘ex larva' 27.V.2005; Luka, 25.V.2007, reared 1♂.

Orius niger is widespread in Western Palaearctic, very rare in the Mediterranean region, also reported from China. It is a very polyphagous species, preying on aphids, psyllids, whiteflies, thrips, larvae of noctuids, mites (Péricart 1972, Herard 1986, Protić 1993).

In south France, Herard (1986) determined Orius niger on pears as a predator of Cacopsylla pyri, but also collected it in a large number from Trioza urticae from nettle, which was surrounding the pear orchards. In Croatia, Orius niger was determined along with Orius minutus on Panonychus ulmi in an apple orchard (Arčanin and Balarin 1972).

In Serbia, Grbić et al. (1989), investigating pear psyllid predators, reported Orius spp., so we are not able to compare our results with theirs. In the same paper, authors mentioned the presence of other Orius species frequently during summer and autumn which is in accordance with our investigations.

Miridae
8) Atractotomus mali Meyer-Dür, 1843
Trophic status.

Phytozoophagous.

Distribution.

Palaearctic.

Preys.

Cacopsylla affinis, from Crataegus monogyna, Koruška, 1.V.2008, reared 1♀ ‘ex larva' 8.V.2008. Cacopsylla melanoneura, from Crataegus monogyna, Beograd–Hotel Jugoslavija, 22.IV.2008, reared 2♂♂, 1♀ ‘ex larva' 30.IV.2008; 18.V.2008, reared 1♀ ‘ex larva' 23.V.2008; Dobra, 10.V.2008, reared 1♂, 1♀ ‘ex larva' 21.V.2008; Draževac, 20.IV.2008, reared 1♂, 1♀ 2.V.2008; Koruška, 1.V.2008, reared 1♀ ‘ex larva' 8.V.2008. Cacopsylla melanoneura, from Malus domestica, Ilinci, 24.V.2008, reared 1♀ ‘ex larva' 30.V.2008; Oparić, 4.V.2008, reared 2♂ ‘ex larva' 19.V.2008; Ilinci, 2.V.2010, reared 1♀. Cacopsylla peregrina, from Crataegus monogyna, Beograd–Hotel Jugoslavija, 22.IV.2008, reared 2♂♂, 1♀ ‘ex larva' 30.IV.2008; Dobra, 10.V.2008, reared 1♂, 1♀ ‘ex larva' 21.V.2008; Koruška, 1.V.2008, reared 1♀ ‘ex larva' 8.V.2008. Cacopsylla picta, from Malus domestica, Ilinci, 24.V.2008, 1♀ ‘ex larva' 30.V.2008.

Atractotomus mali has been reported so far as a predator of mites, aphids, thrips, psyllids, butterfly larvae and pupae (Wheeler 2000b). It was registered as a predator of apple psyllid Cacopsylla mali in Norway (Jonsson 1983) and of pear psyllid Cacopsylla pyrisuga in Greece (Santas 1987). In Germany, Novak and Achtziger (1995) registered it as a predator of hawthorn psyllids Cacopsylla spp.

First record of Atractotomus mali as a predator of Cacopsylla picta. The above mentioned data are the first ones for Atractotomus mali as a psyllid predator in Serbia.

9) Campylomma verbasci (Meyer-Dür, 1843)
Trophic status.

Zoophytophagous.

Distribution.

Holarctic.

Preys.

Cacopsylla bidens, from Pyrus communis, Ilinci, 24.V.2008, reared 1♂ ‘ex larva' 29.V.2008; Bavanište, 25.V.2006, reared 1♂, 1♀ ‘ex larva' 30.V.2006. Cacopsylla pyri, Cacopsylla pyricola and Cacopsylla pyrisuga, from Pyrus communis, Bavanište, 25.V.2006, 1♂, 1♀ ‘ex larva' 30.V. 2006. Cacopsylla melanoneura, from Malus domestica, Ilinci, 2.V.2010, reared 2♂♂.

Campylomma verbasci is a zoophytophagous species preying on apple aphids, pear psyllids, codling moth, thrips and mites (Wheeler 2000b). Its most common prey among insects are Aphis pomi and Cacopsylla mali, and among mites Panonychus ulmi and Tetranychus urticae (Hagen et al. 1999, Wheeler 2000b, Bradley 2007).

However, if there is a lack of prey, it can feed on apple fruits, rarely pear, causing the harm to their aesthetic value. Therefore, Campylomma verbasci is a significant fruit pest in Canada (Hagen et al. 1999, Wheeler 2000a, Bradley 2007). Erler (2004) reported the presence of Campylomma verbasci as a predator of Cacopsylla pyrisuga in treated and untreated pear orchards in Turkey, and Harizanova et al. (2012) mentioned it on Acizzia jamatonica in Bulgaria.

Already known in Serbia (Protić 1993) but in our investigations registered for the first time as a predator of psyllids in this country.

10) Deraeocoris (Deraeocoris) flavilinea (A. Costa, 1862)
Trophic status.

Zoophytophagous.

Distribution.

Western and Central Europe.

Preys.

Cacopsylla bidens, from Pyrus communis, Beograd–Karaburma, 4.V.2006, reared 1♀ ‘ex larva' 18.V.2006. Homotoma ficus, from Ficus carica, Beograd–Banjica, 21.V.2009, collected 1♂, 2♀♀. Trioza rhamni, from Rhamnus cathartica, Beograd–Hotel Jugoslavija, 26.V.2005, reared 1♂ ‘ex larva' 29.V.2005. Psyllopsis discrepans and Psyllopsis machinosa, from Fraxinus angustifolia, Beograd–Autokomanda, 21.V.2009, 1♂, 1♀.

Deraeocoris flavilinea is reported so far as a predator of psyllids (Jerinić-Prodanović and Protić 2011, Simov et al. 2012). Until 1980’s, it was known only from Sicily, from where it has spread to Central Europe where it is now considered as an invasive species (Rabitsch 2008). As a predator of psyllids, it has been known in Serbia since 2011 (Jerinić-Prodanović and Protić 2011).

11) Deraeocoris (Deraeocoris) ruber (Linnaeus, 1758)
Trophic status.

Zoophytophagous.

Distribution.

Holarctic.

Prey.

Cacopsylla pyri, from Pyrus communis, Radmilovac, 10.VII.2006, reared 1♀.

A very polyphagous zoophytophagous species. A Holarctic species occurring in large quantities in the south of Europe.

Already mentioned as a predator of Cacopsylla pyrisuga (Herard 1986). It also preys on younger caterpillar instars of some butterflies, mites and various other small insects in apple orchards, on Rubus spp. and Urtica spp. as well as on aphids on Corylus spp. (Herard 1986).

Reported as a predator of Acizzia jamatonica (Harizanova et al. 2012) in Bulgaria and Cacopsylla pyrisuga in Serbia (Jerinić-Prodanović et al. 2010).

12) Deraeocoris (Knightocapsus) lutescens (Schilling, 1836)
Trophic status.

Zoophagous.

Distribution.

Euro-Mediterranean.

Preys.

Cacopsylla pyri, from Pyrus communis, Izvor, 14.IV.2009, 1♂. Cacopsylla rhamnicola, from Rhamnus cathartica, Obedska bara, 4.VI.2005, reared 1♂ ‘ex larva' 18.VI.2005. Camarotoscena speciosa, from Populus nigra, Zemun–Nova Galenika, 18.X.2010, collected 1♀. Psyllopsis fraxinicola, from Fraxinus angustifolia, Zemun–Nova Galenika, 1.IX.2006, collected 1♀. Trioza chenopodii, from Atriplex oblongifolia, Galovica, 18.VII.2003, reared 1♂.

Deraeocoris lutescens is a Mediterranean species, distributed also in small numbers in Central Europe. Known mainly as an egg predator of pear psyllid Cacopsylla pyrisuga and hawthorn psyllid Cacopsylla crataegi (Herard 1986). It is also reported as a predator of aphids and mite Panonychus ulmi in apple orchards in Croatia (Arčanin and Balarin 1972) and in pear orchards as a predator of Cacopsylla pyrisuga in France and Turkey (Herard 1986, Erler 2004).

Deraeocoris lutescens has been already registered in Serbia (Protić 1993) but here is reported for the first time as a predator of psyllids in this country.

13) Heterocordylus (Heterodactylus) genistae (Scopoli, 1763)
Trophic status.

Phytozoophagous.

Distribution.

Europe.

Prey.

Cacopsylla melanoneura, from Malus domestica, Beloljin, 4.V.2008, collected 1♂; Ilinci, 20.V.2006, reared 1♂ ‘ex larva' 25.V.2006; Uzovnica, 29.IV.2007, collected 1 specimen.

Heterocordylus genistae is mentioned in the literature as a beneficial insect being a predator of psyllids both in larval and adult stage. It is registered as a predator of various other insects (Protić 1993, 1998).

In the present paper, we report Heterocordylus genistae for the first time as a predator of psyllids in Serbia.

14) Hypseloecus visci (Puton, 1888)
Trophic status.

Zoophagous.

Distribution.

Europe.

Prey.

Cacopsylla visci, from Viscum album, Beograd–Bulevar Aleksandra Karađorđevića, 25.III.2007, reared 4♀♀ ‘ex larva' 16.IV.2007.

An exclusively zoophagous species.

Already known from Serbia as a psyllid predator (Jerinić-Prodanović and Protić 2011).

15) Malacocoris chlorizans Panzer, 1794
Trophic status.

Zoophagous.

Distribution.

Eurasia.

Prey.

Homotoma ficus, from Ficus carica, Beograd–Hotel Jugoslavija, 16.V.2007, collected 1♂, 1♀; Beograd–Voždovac, 26.V.2007, collected 5♂♂, 1♀; Zemunski kej, 15.V.2008, collected 1 larva.

A general predator on aphids, psyllids, eggs and larvae of leaf miner moths (Wheeler 2000b, Wyniger and Burckhardt 2003). In Croatia, it is registered as a predator of Panonychus ulmi in apple orchards by Arčanin and Balarin (1972).

Malacocoris chlorizans has been already registered in Serbia (Protić 1998) but in the present paper is reported for the first time as a predator of psyllids in this country.

16) Miris striatus (Linnaeus, 1758)
Trophic status.

Zoophagous.

Distribution.

Europe, Central Asia.

Preys.

Cacopsylla melanoneura and Cacopsylla peregrina, from Crataegus monogyna, Dobra, 10.V.2008, reared 1♀ ‘ex larva' 21.V.2008.

Already reported from Serbia (Protić 1993, 1998). The above mentioned record is the first one of Miris striatus as a predator of psyllids.

17) Orthotylus (Orthotylus) marginalis Reuter, 1884
Trophic status.

Zoophytophagous.

Distribution.

Eurosiberian.

Prey.

Cacopsylla rhamnicola, from Rhamnus cathartica, Beograd–Hotel Jugoslavija, 15.V.2008, reared 1♀ ‘ex larva' 19.V.2008.

Orthotylus marginalis is registered as a predator of aphids and psyllids (Wheeler 2000b). In Finland and Russia, it was mentioned as a predator of Cacopsylla mali (Jonsson 1983).

Registered in Serbia (Protić 2011) but here reported for the first time as a psyllid predator.

18) Psallus (Hylopsallus) assimilis Stichel, 1956
Trophic status.

Phytozoophagous.

Distribution.

Europe.

Preys.

Psyllopsis discrepans, from Fraxinus sp., Ilinci, 21.V.2005, reared 2♂♂, 2♀♀; Umčari, 25.V.2007, reared 1♀. Psyllopsis fraxinicola and Psyllopsis discrepans, from Fraxinus sp., Beograd–Autokomanda, 10.V.2007, reared 5♂♂, 8♀♀ ‘ex larva' 16.V.2007.

Already known as a predator of various insect species, inlcuding psyllids.

Previously registered in Serbia (Protić 1998) but in the present paper reported for the first time as a psyllid predator in this country.

19) Psallus (Psallus) flavellus Stichel, 1933 Fig. 3
Trophic status.

Phytozoophagous.

Distribution.

Europe.

Preys.

Psyllopsis spp., from Fraxinus sp., Beograd–Autokomanda, 8.V.2010, reared 3♂♂, 1♀. Psyllopsis discrepans, from Fraxinus sp., Beograd–Autokomanda, 13.V.2010, reared 2♂♂, 1♀.

Previously registered in Serbia (Protić 2011). Reported here for the first time as a psyllid predator.

Figure 3.

Larva of Psallus flavellus feeding on Psyllopsis fraxinicola.

20) Psallus (Phylidea) quercus Kirschbaum, 1856
Trophic status.

Phytozoophagous.

Distribution.

Europe, Asia.

Preys.

Psyllopsis discrepans, from Fraxinus ornus, Ilinci, 24.V. 2008, collected 1♀; 17.V. 2009, 1♂, 2♀♀. Psyllopsis discrepans and Psyllopsis machinosa, from Fraxinus sp., Beograd–Autokomanda, 6.V.2009, reared 4♀♀ ‘ex larva' 10.V.2009; 21.V.2009, reared 1♂, 3♀♀.

So far known as a predator of aphids, psyllids, thrips, spiders and eggs of various insects (Protić 1998).

Registered in Serbia (Protić 2011) but here reported for the first time as a psyllid predator in this country.

21) Pseudoloxops coccinea Meyer-Dür, 1843
Trophic status.

Zoophagous.

Distribution.

Euro-Mediterranean.

Preys.

Psyllopsis fraxinicola, from Fraxinus sp., Niš, 27.V.2008, collected 1♂. Psyllopsis sp., Fraxinus sp., Beograd–Autokomanda 8.V.2010, reared 1♂ ‘ex larva' 21.V.2010.

Registered in Serbia (Protić 1999). Reported for the first time as a predator of psyllids.

Conclusions

From 28 psyllid species and 19 host plants, we reared or collected 21 species of true bugs belonging to the families Anthocoridae and Miridae. According to available literature data, 12 of the recorded species are zoophagous, while the other nine have mixed nutrition.

Miris striatus, Pseudoloxops coccinea and Psallus flavellus (Miridae) have not been registered as psyllid predators so far. Sixteen species of true bugs are recorded here for the first time as psyllid predators in Serbia (Anthocoris amplicollis, Anthocoris nemorum, Anthocoris confusus, Orius majusculus, Orius niger, Atractotomus mali, Campylomma verbasci, Deraeocoris lutescens, Heterocordylus genistae, Malacocoris chlorizans, Orthotylus marginalis, Psallus assimilis, Psallus quercus, Psallus flavellus, Miris striatus and Pseudoloxops coccinea).

From the family Anthocoridae, we identified seven species: Anthocoris amplicollis, Anthocoris confusus, Anthocoris nemoralis, Anthocoris nemorum, Orius majusculus, Orius minutus and Orius niger. The most polyphagous among them was Orius minutus, found on 13 species of psyllids: Baeopelma foersteri, Cacopsylla melanoneura, Cacopsylla picta, Cacopsylla pyrisuga, Cacopsylla rhamnicola, Camarotoscena speciosa, Homotoma ficus, Psyllopsis discrepans, Psyllopsis fraxinicola, Psyllopsis machinosa, Psyllopsis repens, Trioza mesembrina and Trioza rhamni.

From the family Miridae, we reared or collected 14 species: Atractotomus mali, Campylomma verbasci, Deraeocoris flavilinea, Deraeocoris ruber, Deraeocoris lutescens, Heterocordylus genistae, Hypseloecus visci, Malacocoris chlorizans, Miris striatus, Orthotylus marginalis, Psallus assimilis, Psallus flavellus, Psallus quercus and Pseudoloxops coccinea. Among them, the most polyphagous were Campylomma verbasci, Deraeocoris flavillinea and Deraeocoris lutescens, each registered on five psyllid species.

Most of predatory true bugs are registered on deciduous perennial plants. We found the highest number of predatory true bugs on psyllids which overwinter on host plant and have more than one generation per year, e.g. Cacopsylla pyri, Psyllopsis fraxinicola and Psyllopsis discrepans. On each of them, seven predatory true bugs were registered. Species from the genus Psallus were registered as predators only of psyllid genus Psyllopsis.

Further investigations are necessary for the preservation of known beneficial predatory true bugs and finding of new ones, potentially usable for biological control on economically significant psyllid species.

Acknowledgments

We thank Alexi Popov and two anonymous reviewers for comments and suggestions on the manuscript. They helped us improve our manuscript. We owe special thanks also to Goran Prodanović and Dragica Smiljanić, who collected a part of insect material.

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