Potential of astigmatid mites (Acari: Astigmatina)
as prey for rearing edaphic predatory mites
of the families Laelapidae and Rhodacaridae (Acari:
Mesostigmata)
Marina F. C. Barbosa1•Gilberto J. de Moraes1
Received: 22 January 2016 / Accepted: 3 April 2016 / Published online: 26 April 2016 Ó Springer International Publishing Switzerland 2016
Abstract Laelapidae and Rhodacaridae are important families of edaphic predatory mites and species of these families have been considered for use in biological control programs of soil pests. Mites of Cohort Astigmatina (Acari: Sarcoptiformes) have been largely used as factitious prey in the mass rearing of various edaphic or plant-inhabiting predatory mites. Stratiolaelaps scimitus (Womersley) (Mesostigmata: Laelapidae) (widely commercialized for the control of fungus gnats and thrips) and Protogamasellopsis zaheri Abo-Shnaf, Castilho and Moraes (Mesostigmata: Rhodacaridae) (not available commercially but promising for the control of thrips and nematodes) are known to be reared on Tyrophagus putrescentiae (Schrank) (Astigmatina: Acaridae), but the possibility to find a perhaps more efficient prey has not been evaluated. The objective of this paper was to evaluate different astigmatid species as prey for these predators. S. scimitus and P. zaheri oviposited on all evaluated astigmatids and the acarid mites T. putrescentiae and Aleuroglyphus ovatus (Tropeau) were the most suit-able prey; to confirm the effect of prey on oviposition rates, pregnant females of the predators were kept under starvation conditions and oviposition was negligible or null. Survivorship was always higher than 78 % and was not influenced by prey species or starvation.
Keywords Augmentative biological control Mass rearing Astigmatina Laelapidae Rhodacaridae
Introduction
The literature about the ability of predatory mites of the order Mesostigmata to consume small arthropods commonly found in agricultural areas is extensive, indicating the role played by those organisms in nature and their potential to be used in applied biological control, as extensively discussed in various chapters of Carrillo et al. (2015).
& Marina F. C. Barbosa mferrazb@yahoo.com.br
1
Depto. Entomologia e Acarologia, ESALQ-USP, Piracicaba, Sa˜o Paulo 13418-900, Brazil DOI 10.1007/s10493-016-0043-4
Biological control of mite and small insect pests have been extensively used in several countries around the world. While the control of plant-inhabiting pests has been done mostly with the use of mesostigmatid predators of the family Phytoseiidae (Sabelis and van Rijn1997; Gerson et al.2003; McMurtry et al.2015), edaphic pests have been controlled with mesostigmatid predators of other families, especially Laelapidae (Navarro-Campos et al. 2012; Colloff et al. 2013; Wu et al.2014; Moreira and Moraes 2015; Saito and Brownbridge 2016). Potential for using mesostigmatids of other families also exists, including those of the family Rhodacaridae (see revision in Castilho et al.2015).
Laelapid predators of the genus Stratiolaelaps are commercialized by at least 11 com-panies in the Americas and Europe (Moreira and Moraes2015) for the control of species of fungus gnats (Bradysia spp.; Diptera: Sciaridae), thrips (Frankliniella occidentalis (Per-gande), Thrips spp. and Echinothrips americanus Morgan; Thysanoptera: Thripidae) and a shore fly (Scatella stagnalis (Fallen); Diptera: Ephydridae). Various companies commer-cialize Stratiolaelaps species under two different names, Stratiolaelaps scimitus (Womers-ley) and Stratiolaelaps miles (Berlese), the second by more companies than the first (Moreira and Moraes2015). Walter and Campbell (2003) concluded that S. miles, considered a cos-mopolitan species, actually is a complex of cryptic species. Cabrera et al. (2005) demon-strated that several commercial populations of S. miles, including those used on studies performed by Chambers et al. (1993), Wright and Chambers (1994) and Ydergaard et al. (1997), actually correspond to S. scimitus, indicating that this may be the only species commercialized. Other species of this family, Gaeolaelaps aculeifer (Canestrini), is com-mercialized by two companies in Europe and Australia (Moreira and Moraes2015).
The rhodacarid Protogamasellopsis zaheri Abo-Shnaf, Castilho and Moraes is not available commercially, but Castilho et al. (2009) demonstrated its ability (mentioned as Protogamasellopsis posnaniensis Wisniewski and Hirschmann, according to R. de C. Castilho, personal communication) to reproduce on various prey, namely Bradysia matogrossensis (Lane), F. occidentalis, the acarid mites Tyrophagus putrescentiae (Schrank) and Rhizoglyphus echinopus (Fumouze and Robin), and the bacteriophagous nematode Rhabditella axei (Cobbold) (mentioned as Protorhabditis sp., according to R. de C. Castilho, personal communication). Among these prey, highest oviposition and/or prey consumption rates were observed on T. putrescentiae, followed by F. occidentalis and R. axei (Castilho et al.2009).
Although extensive use of a few species of predatory mites has also been done in open fields, especially in orchards (Koehler1997; Navarro-Campos et al.2012; Colloff et al.2013; Schmidt et al.2013; Tixier et al.2014; Childers and Ueckermann2015), the practical use of most predatory mites for pest control has been done mostly on protected crops (Chambers et al.1993; Lesna et al.1996; Freire et al.2007; Zhang2003; Castilho et al.2009; Gerson and Weintraub2012; McMurtry et al.2013). Cash crops are heavily damaged by most of the same mites that also attacks protected crops. However, biological control of pest mites on the former is hampered by the much larger area cultivated by each grower and the concurrent comparatively lower revenue per unit area. Most certainly considerable efforts have been dedicated by private companies to develop less costly production techniques, although for obvious reasons this information is not published by these companies.
Methods for rearing S. scimitus under laboratory conditions were reported by Wright and Chambers (1994), Steiner et al. (1999) and Cabrera et al. (2005). A pilot system for the mass production of S. scimitus was proposed by Freire and Moraes (2007).
The ability of predatory mites to survive and reproduce when fed with astigmatids is very auspicious, given that several of these mites are easily produced in large numbers on
Hughes1976). This in turn leads to reduced production cost (Gerson et al.2003), making biological control more competitive with other control methods.
Although it is known that both S. scimitus and P. zaheri may reach high population levels when fed with T. putrescentiae (Freire and Moraes2007; Castilho et al.2009), the possibility to use other Astigmatina as prey, perhaps more efficiently, has not been eval-uated. The objective of this paper was to evaluate different species of astigmatid mites as food sources for those predators.
Materials and methods
Stock colonies and their maintenance
The 11 astigmatid species evaluated as prey were Acalvolia squamata (Oudemans) (Winterschmidtiidae), Aeroglyphus robustus (Banks) (Aeroglyphidae), Aleuroglyphus ovatus (Tropeau), Cosmoglyphus oudemansi (Zachvatkin), Thyreophagus cracentiseta Barbosa, OConnor and Moraes and T. putrescentiae (Acaridae), Blomia tropicalis Bron-swijk, de Cock and Oshima (Echimyopodidae), Chortoglyphus arcuatus (Tropeau) (Chortoglyphidae), Dermatophagoides pteronyssinus (Trouessart) (Pyroglyphidae), Gly-cyphagus domesticus (De Geer) (Glycyphagidae) and Suidasia nesbitti Hughes (Suidasi-idae). T. putrescentiae was evaluated only for S. scimitus, because although known to be a suitable prey for this predator, its oviposition capacity on T. putrescentiae was never quantified (quantified for P. zaheri by Castilho et al.2009). Predators were obtained from colonies maintained at Escola Superior de Agricultura ‘‘Luiz de Queiroz’’ (Esalq), Piracicaba, Sa˜o Paulo, Brazil, fed with a mixture of all stages of T. putrescentiae.
Stock colonies of astigmatids and predators were reared in plastic containers similarly to those described by Freire and Moraes (2007), consisting of plastic pots (12 cm high, 7.5 cm diameter), each containing holes for ventilation (2 cm diameter) closed with a polyester screen of 0.2 mm mesh. They were maintained at 25 ± 1°C, 75 ± 10 % rela-tive humidity and in the dark and the astigmatids were fed with 50 % of brewer’s yeast and 50 % of wheat germen. Rearing C. oudemansi using this system lead to the death of mites due to dryness, so this species was reared in a plastic container with the bottom covered with a layer of a solidified paste made of a mixture (9v:1v) of gypsum and activated charcoal kept humid by daily addition of water (Abbatiello 1965) and fed on humidified brewer´s yeast.
Oviposition test
Experimental units were maintained in incubators at 25 ± 1°C, 90 ± 10 % RH and in the dark. Each experimental unit consisted of a plastic dish (2.7 cm diameter, 1.2 cm high) about half filled with the solidified paste of gypsum and activated charcoal, with humidity maintained by daily additions of distilled water. Each unit was sealed with a piece of transparent plastic film (MagipackÒ) to prevent mites from escaping. Predators were transferred to new experimental units every other day, to avoid possible negative effect of accumulated debris and, mainly, to facilitate the detection of the eggs laid.
A visually healthy and gravid predator female (n = 30) was transferred from the stock colony to an experimental unit and fed ad libitum with a mixture of all stages of the evaluated prey. Each experimental unit was examined daily for 11 consecutive days, to
determine the rates of oviposition and survivorship, as well as to replace the available food. Other 30 similar predator females of each species were isolated in experimental units without food as controls. Eggs laid on the first day were excluded from analysis to prevent possible influence on the results.
Data analysis
Statistical analysis were performed on SAS University Edition. Because the data did not satisfy the assumptions of normality (Shapiro Wilk’s test) and homoscedasticity (Levene test), nonparametric tests were used to determine statistical significance (Kruskal–Wallis ANOVA) and to compare treatments (Mann–Whitney U test). Survivorship was analyzed using the Chi square tests.
Results
Both predators oviposited on all evaluated prey. For both predators, significant differences were observed between oviposition levels on different prey. (Table1; S. scimitus: H = 200.14, df = 11; P. zaheri: H = 204.44, df = 10, both p \ 0.0001).
Highest oviposition of S. scimitus was obtained when fed on T. putrescentiae, followed by A. ovatus. Lowest oviposition rates were obtained when fed on A. robustus and C. arcuatus. Oviposition was negligible (0.1 egg/ female/ day) when no food was offered. Survivorship of the predator did not vary with prey species and was always at least 79 % (v2= 7.81, df = 11, p = 0.64) at the end of the study (Table1).
Table 1 Mean (±SE) daily oviposition and survival (after 11 days) of Stratiolaelaps scimitus and Pro-togamasellopsis zaheri fed different astigmatid prey species at 25 ± 1°C, 70 ± 10 % RH and in the dark
S. scimitus P. zaheri n Daily oviposition Survival (%) n Daily oviposition Survival (%) Tyrophagus putrescentiae 29 3.4 ± 0.10 a 97 a –a Aleuroglyphus ovatus 29 2.3 ± 0.10 b 93 a 26 8.5 ± 0.40 a 92 a Suidasia nesbitti 29 1.7 ± 0.09 c 97 a 25 3.6 ± 0.12 de 100 a Acalvolia squamata 30 1.6 ± 0.08 cd 90 a 26 4.5 ± 0.16 c 88 a Cosmoglyphus oudemansi 28 1.5 ± 0.11 cd 93 a 26 5.1 ± 0.13 b 92 a Glycyphagus domesticus 28 1.5 ± 0.07 cd 93 a 28 2.5 ± 0.13 f 93 a Thyreophagus cracentiseta 29 1.4 ± 0.08 d 79 a 23 3.8 ± 0.29 de 78 a Dermatophagoides pteronyssinus 29 1.2 ± 0.08 de 90 a 28 2.0 ± 0.11 g 93 a Blomia tropicalis 30 1.0 ± 0.12 e 90 a 25 3.8 ± 0.14 d 84 a Aeroglyphus robustus 27 0.6 ± 0.03 f 93 a 27 3.2 ± 0.18 e 93 a Chortoglyphus arcuatus 29 0.6 ± 0.08 f 90 a 25 2.2 ± 0.09 fg 92 a Without food 28 0.1 ± 0.01 g 96 a 29 0 h 93 a Means within a column followed by the same letter are not significantly different (daily oviposition: Kruskal–Wallis ANOVA, Mann–Whitney U test; survival: Chi square test; p \ 0.05)
a
Highest oviposition of P. zaheri was obtained when fed on A. ovatus, followed by C. oudemansi. Lowest oviposition rate was obtained when fed on C. arcuatus. No eggs were laid in the absence of food. Also in this case survivorship of the predator did not vary with prey species and was always at least 78 % (v2= 9.44, df = 10, p = 0.49) (Table1).
Discussion
The much higher oviposition rates of P. zaheri on all tested prey compared with those of S. scimitus was already expected, given the results of previous studies offering other food types to those species (Ali and Brennan1997; Enkegaard et al.1997; Cabrera et al.2005; Castilho et al.2009). Interestingly, the two prey allowing the highest oviposition rates in this study for S. scimitus (A. ovatus and T. putrescentiae) were also the ones allowing the highest oviposition rates for P. zaheri, as indicated by the results of this study comple-mented by the results of the study of Castilho et al. (2009).
Daily oviposition rates determined in this study for S. scimitus fed on A. ovatus and T. putrescentiae were high compared with data of previous studies: 1.4 on Bradysia sp. larvae and 2.0 on unidentified Enchytraeidae (Oligochaeta) (Cabrera et al. 2005). When fed on Sancassania aff. sphaerogaster, immature of S. scimitus did not reach the deutonymphal stage (Cabrera et al.2005).
Oviposition rates of S. scimitus comparable to the best values obtained in this study was reported by Ali and Brennan (1997) on the acarid Acarus siro L. (2.6 eggs/female/day) and much lower levels were reported by Shereef et al. (1980) and Enkegaard et al. (1997) on T. putrescentiae (0.7) and by Shereef et al. (1980) and Hoda et al. (1986) on R. echinopus (0.8); in these papers, the mite species was named as S. miles, but considering the findings in Cabrera et al. (2005), they may actually be S. scimitus.
Oviposition of P. zaheri fed on A. ovatus, was slightly higher than reported by Castilho et al. (2009) on T. putrescentiae (7.6 eggs/female/day) and much higher than reported by the same authors on R. echinopus (1.9).
Cosmoglyphus oudemansi also promoted high oviposition of P. zaheri in this study, but the mass rearing of that prey seems more difficult than the rearing of other prey, due to its requirement for levels of relative humidity nearing saturation (our personal observation). Aleuroglyphus ovatus and T. putrescentiae, ubiquitous acarids (Oconnor 2009), can be easily reared on several cheap substrates (e.g. Rivard1961; Sasa et al.1970; Chmielewski
1999; Canfield and Wrenn2010; Xia et al.2009), what is an additional advantage of using these species in the mass production of those and possibly other predators of the same families.
Castilho et al. (2009) assumed that the ability of P. zaheri to feed on astigmatids suggested it to be a generalist predator, given that astigmatids are often uncommon in soils, habitat where those predators are regularly found in Brazil. The same argument could apply to S. scimitus, which is also found in Brazil in microhabitats where astigmatids are uncommon. The acceptance of all evaluated prey as food by these two predator in the present study corroborates the assumption of Castilho et al. (2009).
High survivorship of P. zaheri and Stratiolaelaps species was already reported on different prey (Enkegaard et al. 1997; Castilho et al. 2009) and in the absence of food (Ignatowicz1974; Wright and Chambers1994; Moreira and Moraes2015). The ability to stand starvation is an important characteristic to be considered in their practical use for pest control, but the evaluation of survivorship as a factor to determine the suitability of
potential prey for these predators should be seen with reservation. Wright and Chambers (1994) mentioned that the oviposition behavior of S. miles involved the search for par-ticular sites in the experimental unit where the eggs would be less exposed. In this study, we observed that females of S. scimitus deposited their eggs in the clusters formed by the prey or other debris formed by prey food. Differently from what was observed when food was available, starved females were observed moving most of the time in search of prey. The experiment setup of this study did not allow the determination of possible differ-ential effects of the debris resulting from the process adopted in the prey rearing (for example, released volatiles, development of infesting microorganisms, etc.). Mass rearing those predators with the prey determined as promising food sources would require eval-uation of those possible effects.
Conclusion
The results obtained, complemented by the results of Castilho et al. (2009), demonstrate the possibility of using A. ovatus as new suitable species for mass rearing S. scimitus and P. zaheri, besides confirming the efficacy of T. putrescentiae as a factitious prey for those predators.
Acknowledgments This work was supported by the Brazilian ‘‘Coordenac¸a˜o de Aperfeic¸oamento de Pessoal de Nı´vel Superior’’ (CAPES) and ‘‘Conselho Nacional de Desenvolvimento Cientı´fico e Tec-nolo´gico’’ (CNPq).
References
Abbatiello MJ (1965) A culture chamber for rearing soil mites. Turtox News 43(7):162–164
Ali O, Brennan P (1997) Development, feeding and reproduction of the predatory mite, Hypoaspis miles (Acari: Mesostigmata: Laelapidae) on different types of prey. Syst Appl Acarol 2(1):81–88 Cabrera AR, Cloyd RA, Zaborski ER (2005) Development and reproduction of Stratiolaelaps scimitus
(Acari: Laelapidae) with fungus gnat larvae (Diptera: Sciaridae), potworms (Oligochaeta: Enchy-traeidae) or Sancassania aff. sphaerogaster (Acari: Acaridae) as the sole food source. Exp Appl Acarol 36:71–81
Canfield MS, Wrenn WJ (2010) Tyrophagus putrescentiae mites grown in dog food cultures and the effect mould growth has on mite survival and reproduction. Vet Dermatol 21(1):58–63
Carrillo D, Moraes GJ, Pen˜a JE (2015) Prospects for biological control of plant feeding mites and other harmful organisms. Springer International, Cham
Castilho RC, Moraes GJ, Silva ES, Silva LO (2009) Predation potential and biology of Protogamasellopsis posnaniensis Wisniewski & Hirschmann (Acari: Rhodacaridae). Biol Control 48(2):164–167 Castilho RC, Venancio R, Narita JPZ (2015) Mesostigmata as biological control agents, with emphasis on
Rhodacaroidea and Parasitoidea. In: Carrillo D, de Moraes GJ, Pen˜a JE (eds) Prospects for biological control of plant feeding mites and other harmful organisms. Springer International, Cham, pp 1–31 Chambers RJ, Wright EM, Lind RJ (1993) Biological control of glasshouse sciarid flies (Bradysia spp.) with
the predatory mite, Hypoaspis miles on cyclamen and poinsettia. Biocontrol Sci Techn 3:285–293 Childers CC, Ueckermann EA (2015) Non-phytoseiid Mesostigmata within citrus orchards in Florida:
species distribution, relative and seasonal abundance within trees, associated vines and ground cover plants and additional collection records of mites in citrus orchards. Exp Appl Acarol 65:331–357 Chmielewski W (1999) Acceptance of buckwheat grain as a food by Tyrophagus putrescentiae (Schr.)
(Acari: Acaridae). Fagopyrum 16:95–97
Colloff MJ, Elizabeth AL, Cook DC (2013) Natural pest control in citrus as an ecosystem service: inte-grating ecology, economics and management at the farm scale. Biol Control 67(2):170–177 Enkegaard A, Sardar MA, Brødsgaard HF (1997) The predatory mite Hypoaspis miles: biological and
demographic characteristics on two prey species, the mushroom sciarid fly, Lycoriella solani, and the mould mite, Tyrophagus putrescentiae. Entomol Exp Appl 82:135–146
Freire RAP, Moraes GJ (2007) Mass production of the predatory mite Stratiolaelaps scimitus (Womersley) (Acari: Laelapidae). Syst Appl Acarol 12(2):117–119
Freire RAP, Moraes GJ, Silva ES, Vaz AC, Castilho RC (2007) Biological control of Bradysia matogrossensis (Diptera: Sciaridae) in mushroom cultivation with predatory mites. Exp Appl Acarol 42:87–93
Gerson U, Weintraub PG (2012) Mites (Acari) as a factor in greenhouse management. Annu Rev Entomol 57:229–247
Gerson U, Smiley RL, Ochoa R (2003) Mites (Acari) for pest control. Blackwell Science, Oxford Griffiths DA (1964) A revision of the genus Acarus L., 1758 (Acaridae, Acarina). Bull Br Mus Nat Hist
(Zool) 11:415–464
Hoda FM, Taha HA, Ibrahim GA, El-Beheri MM (1986) Biological observations on the predator mite, Hypoaspis miles Berlese (Acarina: Laelapidae). Bull Soc Entomol Egypte 66:103–106
Hughes AM (1976) The mites of stored food and houses Tech. Bull. No. 9. Ministry of Agriculture, Fisheries and Food, London
Ignatowicz S (1974) Observations on the biology and development of Hypoaspis aculeifer Canestrini, 1885 (Acarina, Gamasides). Zool Pol 24:41–59
Koehler HH (1997) Mesostigmata (Gamasina, Uropodina), efficient predators in agroecosystems. Agr Ecosyst Environ 62:105–117
Lesna I, Sabelis M, Conijn C (1996) Biological control of the bulb mite, Rhizoglyphus robini, by the predatory mite, Hypoaspis aculeifer, on lilies: predator-prey interactions at various spatial scales. J Appl Ecol 33(2):369–376
Mcmurtry JA, Moraes GJ, Sourassou NF (2013) Revision of the lifestyles of phytoseiid mites (Acari: Phytoseiidae) and implications for biological control strategies. Syst Appl Acarol 18(4):297–320 McMurtry JA, Sourassou NF, Demite PR (2015) The Phytoseiidae (Acari: Mesostigmata) as biological
control agents. In: Carrillo D, de Moraes GJ, Pen˜a JE (eds) Prospects for biological control of plant feeding mites and other harmful organisms. Springer International, Cham, pp 133–149
Moreira GF, Moraes GJ (2015) The potential of free-living Laelapid mites (Mesostigmata: Laelapidae) as biological control agents. In: Carrillo D, de Moraes GJ, Pen˜a JE (eds) Prospects for biological control of plant feeding mites and other harmful organisms. Springer International, Cham, pp 77–102 Navarro-Campos C, Pekas A, Moraza ML, Aguilar A, Garcia-Marı´ F (2012) Soil-dwelling predatory mites
in citrus: their potential as natural enemies of thrips with special reference to Pezothrips kellyanus (Thysanoptera: Thripidae). Biol Control 63:201–209
OConnor BM (2009) Cohort Astigmatina. In: Krantz GW, Walter DE (eds) A manual of Acarology. Texas Tech University Press, Austin, pp 565–657
Rivard I (1961) Influence of temperature and humidity on longevity, fecundity, and rate of increase of the mite Tyrophagus putrescentiae (Schrank) (Acarina: Acaridae) reared on mold cultures. Can J Zool 39:869–876
Sabelis MW, van Rijn PCJ (1997) Predation by insects and mites. In: Lewis T (ed) Thrips as crop pests. CAB-International, London, pp 259–354
Saito T, Brownbridge M (2016) Compatibility of soil-dwelling predators and microbial agents and their efficacy in controlling soil-dwelling stages of western flower thrips Frankliniella occidentalis. Biol Control 92:92–100
Sasa M, Miyamoto J, Shinohara S, Suzuki H, Katsuhata A (1970) Studies on mass culture and isolation of Dermatophagoides farinae and some other mites associated with house dust and stored food. Jpn J Exp Med 40:367–382
Schmidt RA, Beers EH, Unruh TR, Horton DR (2013) Releases of insectary-reared Galendromus occi-dentalis (Acari: Phytoseiidae) in commercial apple orchards. J Econ Entomol 106:1996–2005 Shereef G, Soliman Z, Afifi A (1980) Economic importance of the mite, Hypoaspis miles (Berlese)
(Me-sostigmata: Laelapidae) and its life history. Bull Zool Soc Egypt 30:130–138
Sinha R (1979) Role of Acarina in the stored grain ecosystem. Recent Adv Acarol 1:263–271
Steiner M, Goodwin S, Wellham T (1999) A simplified rearing method for Stratiolaelaps (Hypoaspis) miles (Acari: Laelapidae). IOBC/WPRS Bull 22:241–242
Tixier A, Badel E, Franchel J, Lakhal W, Leblanc-Fournier N, Moulia B, Julien JL (2014) Growth and molecular responses to long-distance stimuli in poplars: bending vs flame wounding. Physiol Plant 150:225–237
Walter DE, Campbell NJH (2003) Exotic vs endemic biocontrol agents: would the real Stratiolaelaps miles (Berlese) (Acari: Mesostigmata: Laelapidae) please stand up? Biol Control 26:253–269
Wright E, Chambers R (1994) The biology of the predatory mite Hypoaspis miles (Acari: Laelapidae), a potential biological control agent of Bradysia paupera (Dipt.: Sciaridae). Entomophaga 39:225–235
Wu S, Gao Y, Xu X, Wang E, Wang Y, Lei Z (2014) Evaluation of Stratiolaelaps scimitus and Neoseiulus barkeri for biological control of thrips on greenhouse cucumbers. Biocontrol Sci Technol 24:1110–1121
Xia B, Luo D, Zou Z, Zhu Z (2009) Effect of temperature on the life cycle of Aleuroglyphus ovatus (Acari: Acaridae) at four constant temperatures. J Stored Prod Res 45:190–194
Ydergaard S, Enkegaard A, Brødsgaard HF (1997) The predatory mite Hypoaspis miles: temperature dependent life table characteristics on a diet of sciarid larvae, Bradysia paupera and B. tritici. Entomol Exp Appl 85:177–187
