Ability of Phytoseiulus longipes to control spider mite pests on tomato in European greenhouses. Maxime Ferrero, Serge Kreiter, Marie-Stéphane Tixier

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Maxime Ferrero, Serge Kreiter, Marie-Stéphane Tixier. Ability of Phytoseiulus longipes to control spider mite pests on tomato in European greenhouses.. 6. European Congress of Acarology, European Association of Acarologists (EURAAC). INT., Jul 2008, Montpellier, France. ￿hal-01137131￿

HAL Id: hal-01137131 https://hal.archives-ouvertes.fr/hal-01137131 Submitted on 30 Mar 2015

HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Integrative Acarology Integrative Acarology. Proceedings of the 6th European Congress M. Bertrand, S. Kreiter, K.D. McCoy, A. Migeon, M. Navajas, M.‐S. Tixier, L. Vial (Eds.) European Association of Acarolgists, 2008

ABILITY OF PHYTOSEIULUS LONGIPES TO CONTROL SPIDER MITE PESTS ON TOMATO IN EUROPEAN GREENHOUSES

M. Ferrero, S. Kreiter and M.-S. Tixier

Montpellier SupAgro, UMR CBGP 1062, bât 16, laboratoire d'Acarologie, 2 Place Pierre Viala 34060 Montpellier cedex 01, France

Abstract

Even if many studies dealt with the biological control of spider mites on tomato in greenhouses, no efficient solution has still been found beyond chemicals to get rid of those phytophagous mites. Among them, two species, Tetranychus evansi and T. urticae, can be considered as very serious pests, leading to great damages in tomato crops in Southern Europe. Preliminary experiments showed that Phytoseiulus longipes is a very promising predator of these two species. In the present study, life tables of a Chilean strain of this predator have been calculated, at 25 °C, 80 ± 10 % RH and 16/ 8 (L/ D), in several prey/ plant conditions: P. longipes feeding on T. evansi on tomato, P. longipes feeding on T. urticae on tomato and P. longipes feeding on T. urticae on bean. 88.9 % of the predators did not complete their immature phase while feeding on T. evansi, and life tables could not be calculated. However, while feeding on T. urticae, immature survival was 99.8 % and 90.0 % on bean and tomato, respectively. Immature durations of P. longipes fed with T. urticae were not different, being 4.35 and 4.21 days, on bean and tomato, respectively. The intrinsic rate of increase (rm) was 0.368 and 0.116 female/ female/ day, on bean and tomato, respectively. Those results suggest that the Chilean strain of P. longipes would not be able to control neither T. evansi nor T. urticae in tomato crops. However, it seems able to eat and develop on T. urticae in other crops. Another strain of P. longipes, originating from Brazil, is currently being studied and lead to very enthusiastic perspectives to control spider mites on tomato greenhouses, both T. urticae and T. evansi. Furthermore, experiments are being conducted to try to explain the surprising differences in feeding habits and host plants between the two strains of this predaceous mite.

Key‐words

Tetranychus evansi, Tetranychus urticae, life history, biological control, Solanaceae

Introduction Migeon 2005, Castagnoli 2006, Tsagkarakou 2007) and causes severe injuries, especially to Spanish Although many predatory mite species have been tomato crops (Ferragut pers. comm.). studied to control tetranychids, those pests still cause serious damages to several crops, and In order to get rid of T. evansi, many methods have especially to tomato in greenhouses. In Southern been experimented, like pesticides (Blair 1989, Europe, the main problem has been Tetranychus Mabeya et al. 2003), plant resistance development urticae Koch for many years (Zhang 2003). But (Maluf et al. 2001, Resende et al. 2002, Gonçalves since the early nineties, an invasive pest, et al. 2006, Resende et al. 2008), Tetranychus evansi Baker & Pritchard, which seems entomopathogenic fungi (Humber & Moraes 1981, to be originated from South America (Gutierrez & Wekesa et al. 2005, 2006, 2007) and several Etienne 1986) spreads through Southern Europe predators (Sarmento et al. 2004, Ho et al. 2005, (Ferragut & Escudero 1999; Bolland & Vala 2000, Oliveira et al. 2005) including predatory mites

Montpellier ‐ 21‐25 July 2008 461 (Moraes & Lima 1983; Moraes & McMurtry 1985a; or bean): T. evansi / tomato, T. urticae / bean and Moraes & McMurtry 1985b; Moraes et McMurtry T. urticae / tomato. The combination T. evansi / 1986; Escudero et Ferragut 2005; Rosa et al. 2005, bean has not been tested because this plant was Ferrero et al. 2007, Furtado et al. 2007, Koller et al. unsuitable for rearing our strain of T. evansi. 2007). The only enthusiastic perspective up to now Groups of five prey females were placed using a has been pointed out by Furtado et al. (2006) and thin paintbrush in experimental units, consisting of Ferrero et al. (2007), with the discover and the a leaf disk of a plant substrate (2 cm in diameter) study of a Brazilian strain of Phytoseiulus longipes placed underside up onto a moist disk of filter Evans (named P. longipes B thereafter). Following paper, inside a Petri dish (2 cm in diameter, 1 cm those studies, this type I predator (McMurtry & high). One day later, eggs of P. longipes (between 0 Croft 1997), feeding preferentially on mites of the and 6 hours old) obtained from the intermediate subfamily Tetranychinae, seems actually to be stock colony (consisting in 30 to 50 females reared specific of T. evansi (Furtado et al. 2007). More on a leaf of P. vulgaris cv. Contender placed in a recently, a Chilean strain of this species (written P. plastic tray as described above) were transferred longipes C thereafter) has been found (Ragusa to each experimental unit (one egg/ unit); then pers. comm.). In order to use these strains in closed with a transparent plastic film. To maintain biological control for controlling T. evansi, further humidity, distilled water was added on the filter biological studies are required. This paper aims to paper every day. Periodically (once a week on determine the life history of P. longipes C on tomato, twice on bean) P. longipes individuals several plant substrates/ prey conditions. were transferred to new leaves infested with preys as previously reported. Material and Methods Observations were carried out every 12 hours to determine the duration and the survivorship for Species and strains studied each stage. The strain of P. longipes studied was obtained Reproduction from a colony initiated with specimens collected in Chile, fed with T. urticae, in Nogal, Recently emerged adult P. longipes females obtained were transferred to new experimental Los Andes, Valparaiso Region (Ragusa, pers. units. A male taken from the stock colony was then comm.). Mites were fed with all stages of T. added to each unit containing one female, and a urticae, offered on leaves of Phaseolus new one was added for every male that died or vulgaris L. cv. Contender, placed underside up escaped. At least 30 couples were observed. Daily in rearing units constituted of plastic trays (10 observations were conducted to determine female × 15 cm) bordered with water‐saturated fecundity and survivorship. The eggs laid were cotton to avoid mite escapes and to maintain placed daily in a single unit and reared to the turgescence of the leaves. Rearing units adulthood to determine the secondary sex ratio were placed in climatic units at 25 ± 2 °C, 75 ± (female percentage of the studied female cohort offspring). 10 % RH and 16/ 8 [L/ D]. The T. evansi stock colony was initiated with specimens collected Life Table from a tomato screenhouse at Saint‐Jeannet The life table was constructed considering the (Alpes‐Maritimes, 06, France) in October 2007 females of the cohort studied. The net (Migeon pers. comm.), and reared on reproductive rate (Ro), the mean generation time Lycopersicon esculentum Miller in rearing (T), the intrinsic rate of increase (rm), the doubling units similar to those described above. generation time (Dt), and the finite rate of increase (λ) were calculated using the method The T. urticae stock colony initiated with recommended by Birch (1948): specimens collected in Montpellier, France, Ro = Σ (l × m ) was reared on P. vulgaris cv. Contender in a x x greenhouse. T = Σ (x × lx × mx)/ Σ (lx × mx)

Immature development rm = Ln (Ro) / T

Experimentations were performed at 25 ± 2 °C, 75 Dt = Ln (2) / rm ± 10 % RH and 16/ 8 [L/ D]. Three items were λ = exp (rm) tested, each one being characterised by a prey (T. evansi or T. urticae) and a plant substrate (tomato In those equations, x is the age (with 0.5 for the

Montpellier ‐ 21‐25 July 2008 462 day when eggs had been laid), lx, the cumulative Results female survivorship, and mx, the number of female descendants per female at x. Immature development Egg to adult duration ranged from 4.10 to 4.21 Calculation of a corrected rm value was performed days, for the T. evansi/ tomato and T. urticae/ by iteration. The method, aiming to find rm for tomato conditions, respectively (Table 1). The egg which (1 ‐ Σ exp (‐rm × x) × lx × mx) is minimal, was given by Maia et al. (2000). stage was the longest, varying from 1.80 to 1.96 days. The larval stage was the shortest, varying Analysis of variance (ANOVA) and related Tukey from 0.35 to 0.55 days. No statistical analysis could HSD mean comparison tests were performed to be performed for the item T. evansi/ tomato for determine differences between duration of the the protonymphal, deutonymphal and egg to adult immature phases and adult stages between the stages because too many individuals had died different items tested. prematurely (11.1 % of the immatures tested

The rm iteration and statistical analysis were computed with R (R project 2008).

Table 1. Mean duration (± Standard Error) in days of the immature instars of a Chilean strain of Phytoseiulus longipes for several items prey/ plant substrate, number of replicates (N and immature survival rate).

Stage Item

T. evansi/ tomato T. urticae/ bean T. urticae/ tomato F (df1, df2) (α = 0.05)

Egg 1.80 (0.21) a 1.96 (0.20) b 1.83 (0.00) a Ft < F (2, 109) = 10.99; P = 4.48 × 10‐5

Larva 0.44 (0.17) ab 0.55 (0.22) a 0.35 (0.23) b Ft < F (2, 105) = 8.91; P = 2.66 × 10‐4

Protonymph 1.05 (0.35) 0.85 (0.26) a 1.01 (0.26) b Ft < F (2, 88) = 4.39 ; P = 1.51 × 10 ‐2

Deutonymph 1.00 (0.00) 0.99 (0.18) a 1.01 (0.29) a Ft > F (2, 80) = 0.11; P = 8.9 × 10‐1

Egg to adult 4.10 (0.00) 4.36 (0.27) a 4.21 (0.35) a Ft > F (2, 80) = 2.91; P= 6.03 × 10‐2 N 27 41 44 Immature survival (%) 11.1 99.8 90.9

Ft (F from tables) < F(df1, df2) (F calculated) mean that there are differences between the mean durations for a stage (ANOVA, α = 0.5). Durations followed by a different letter for a stage are significantly different (Tukey HSD test, α = 0.5). No statistical analysis could be performed for some stages of the T. evansi / tomato condition because of high mortality at the larval stage.

Table 2. Mean durations (± Standard Errors) in days of adult phases, longevity and ovipositional rates of Phytoseiulus longipes C feeding on Tetranychus urticae on bean or tomato, number of replicates (N).

Stage Plant substrate

Bean Tomato F (df1, df2) (α = 0.05)

‐1 Pre‐oviposition 1.22 (0.93) a 0.95 (0.69) a Ft > F (1, 40) = 1.05; P = 3.11 × 10

‐1 Oviposition 3.74 (1.85) a 2.63 (2.41) a Ft > F (1, 40) = 2.70; P = 1.08 × 10

‐2 Post‐oviposition 0.30 (0.46) a 1.21 (1.64) b Ft < F (1, 40) = 6.10; P = 1.79 × 10

‐4 ‐1 Longevity 8.95 (2.18) a 8.98 (2.62) a Ft > F (1, 51) = 6.00 × 10 ; P = 9.81 × 10 Eggs / female / day 1.34 (0.89) 0.57 (0.48) Total eggs / female 6.77 (5.79) 3.29 (2.78) N 30 23

Ft (F from tables) < F(df1, df2) (F calculated) mean that there are differences between the mean durations for a stage (ANOVA, α = 0.5). Durations followed by a different letter for a stage are significantly different (Tukey HSD test, α = 0.5).

Montpellier ‐ 21‐25 July 2008 463 Table 3. Phytoseiulus longipes C demographic parameters on two substrates fed with Tetranychus urticae at 25 °C, 80 ± 10 % RH and 16/ 8

(L/ D). Net reproductive rate (Ro), mean generation time (T), finite rate of increase (λ), doubling generation time (Dt), estimated and iterated intrinsic rate of increase (rm).

Substrate Demographic parameter

Ro T λ Dt rm (Birch) rm (iterated) Bean 3.92 4.04 1.40 2.05 0.338 0.368 Tomato 1.39 2.91 1.12 6.12 0.113 0.116

P. longipes C to develop and reproduce fed with survived). No significant differences were found tetranychids on tomato, through the analysis of its between the two other conditions tested for the life parameters The second goal was to compare egg to adult and deutonymph durations. For the the biological characteristics of the studied strains three other immature stages (i.e. egg, larva and of this species. Up to 2007, experiments had been protonymph), significant differences were always performed on two other strains of this predator: a found between the items T. urticae/ bean and T. South‐African strain (reported as P. longipes SA urticae/ tomato. thereafter) and a Brazilian strain (Badii & Reproduction McMurtry 1983, 1984; Moraes & McMurtry 1985b; Takahashi & Chant 1992, Badii et al. 1999). It has Adult phase durations and oviposition rates were been reported that the South‐African strain is observed only when P. longipes was feeding on T. unable to control T. evansi in the conditions tested, urticae, on bean or tomato (Table 2), because of even if promising results had been found to control the high mortality of P. longipes when fed with T. Tetranychus pacificus (McGregor) (Badii et al., evansi. The longevity was not significantly different 1999). On the opposite, the results obtained for between the two conditions, varying from 8.95 and the Brazilian strain, discovered in 2005 (Furtado et 8.98 days, on bean and tomato, respectively. The al. 2006), are enthusiastic for the control of T. pre‐oviposition period ranged from 1.22 to 0.95 evansi but also of T. urticae on tomatoes (Ferrero days, oviposition from 3.74 to 2.63 days and post‐ et al. 2007; Furtado et al. 2007. oviposition from 0.30 to 1.21 days, on bean and tomato, respectively. The only significant Concerning the developmental phase of P. longipes difference found between these two conditions C, several points could be pointed out. At first, for was for the post‐oviposition period. The mean the item T. evansi/ tomato, only 3 individuals out oviposition rate and total oviposition were highest of 27 reached the adult stage, and none of these 3 when P. longipes was fed with T. urticae on bean mites survived enough to mate. These conditions than on tomato, being 1.34 eggs/ female/ day and thus seem to be unsuitable for the Chilean strain of 6.77 eggs/ female, respectively. P. longipes, which is a key difference to notice with P. longipes B, which seems specific to T. evansi Life tables (Furtado et al. 2007). For the other conditions Calculated life table parameters are given in Table tested in the present paper (see table IV), egg to 3. Congruently with the previous observations, the adult durations were lower to what has been best features were obtained for the item T. observed previously with 4.7 and 4.8 days while P. urticae/ bean. The intrinsic rate of increase longipes B was feeding on T. evansi and T. urticae, (iterated) was more than three times higher on respectively, on Canavalia ensiformis (L.) (Furtado bean than on tomato, while the doubling time and et al. 2007), 4.9 days while P. longipes B was the net reproductive rate were almost three times feeding on T. evansi on Solanum americanum lower on bean than on tomato. The calculated Miller (Ferrero et al. 2007) and 5.23 days for P. mean generation time was 4.04 and 2.91 days, and longipes SA feeding on T. pacificus on bean (Badii the finite rate of increase 1.40 and 1.12, on bean & McMurtry 1984). The Chilean strain of P. and tomato, respectively. longipes thus seems to develop faster than the other strains feeding on T. urticae. Discussion, conclusions and perspectives The whole immature phase durations are of the same order of magnitude or longer for the other This study has been performed in order to Phytoseiulus species, with 5.5 days for Phytoseiulus determine first the ability of the predaceous mite

Montpellier ‐ 21‐25 July 2008 464 Table 4. Comparison of immature development durations (in days), longevity (in days) and intrinsic rates of increase (female/ female/ days) for several Phytoseiulus species, depending on the prey provided, the plant substrate and climatic conditions.

2007

al.

2007 Ferragut

al. et

& 2005 al. et

2005 et McMurtry al.

& et

Furtado Escudero 2008 Vasconcelos 1984 Silva Badii Reference Ferrero

of

(female/

m r rate day)

increase 0.293 0.32 0.363 0.366 0.193 0.116 0.373 0.273 female/ Intrinsic

in

Longevity days 20.3 29.7 31.1 34.12 44 NA NA 23.3

duration

days

in 4.9 4.8 4.7 5.23 4.8 6.91 4.16 5.5 Immature development

RH,

D RH,

% %

% % L/

RH,

7 12 10

D D 80 12

% ± ± ±

‐ 5

L/ L/

80 83 80 70 D D D ±

12/

conditions

12 12

L/ L/ L/ °C, °C, °C, °C,

60

1 1 2 0.8, 1

12/ 12/ 12 10 12

± ± ± ± ± °C,

12/ 26 14/ 25 RH, 25 RH, 25 25 12/ Climatic 25

substrate

ensiformis vulgaris lunatus

P. S.americanum Solanum americanum Canavalia ensiformis P. C. Plant

species

urticae urticae evansi urticae pacificus evansi urticae

T. T. T. T. T. T. Prey evansi T. Tetranychus

SA B

macropilis persimilis fragariae longipes longipes

P. P. P. P. P. species Phytoseiulus

Montpellier ‐ 21‐25 July 2008 465 Chilean strain of P. longipes. This work showed fragariae Denmark & Schicha feeding on T. urticae results different than those already obtained for on S. americanum (Vasconcelos et al. 2008), 4.8 other strains of this species. Phytoseiulus longipes days for Phytoseiulus macropilis (Banks) feeding on C was, as the SA strain, unable to complete its T. urticae on C. ensiformis (Silva et al. 2005) and development feeding on T. evansi on tomato, 6.91 and 4.16 days for Phytoseiulus persimilis while the B strain could. Those observations lead Athias‐Henriot feeding on T. evansi and T. urticae to the first major conclusion of this work which is on bean (Escudero & Ferragut 2005). Results that P. longipes C would be unable to control T. obtained here on the developmental phase of P. evansi populations in tomato greenhouses. Great longipes C showed that this strain is not able to differences in longevity had been also pointed out develop, thus to control, T. evansi on tomato, but between the Chilean strain and the two others. can develop in the same way on tomato and bean These differences could be due to differences in while feeding on T. urticae. At last, as defined by the setup (photoperiod), or to the strain itself. McMurtry & Croft (1997), type I phytoseiids, Values of r showed that P. longipes C would be belonging to the genus Phytoseiulus, do not need m able to develop and reproduce well feeding on T. food until they reach the protonymphal stage. This urticae on bean, but apparently not enough has also been observed during the present feeding on T. urticae on tomato to settle an experiment for the Chilean strain of P. longipes enthusiastic prognosis for its use in biological Adult phase durations of the Chilean strain of P. control. longipes were calculated only with T. urticae as In this paper, particular attention was paid to the prey. These durations were very short compared to differences between the three strains of P. all the data previously compiled (table IV). longipes already studied: differences in the Longevity was only 8.95 and 8.98 days in our data suitability of plant substrate and in prey items, in for items T. urticae/ bean and T. urticae/ tomato, differences in longevity and oviposition at the respectively, while Ferrero et al. (2007) showed a same conditions. All these observations lead the 20.3 days longevity for P. longipes B and Badii authors to settle a whole study to compare those (1984) a 34.12 days longevity for P. longipes SA. strains in combined conditions. Along with more For the other Phytoseiulus species studied feeding biological studies, other tools like genetic markers on T. urticae at the same temperatures than in the or morphological analysis are presently being used present study, longevity is also variable, from 44.0 to determine and explain some of those and 23.3 days, for P. macropilis and P. fragariae, differences. Many conclusions on the usefulness respectively (Silva et al. 2005; Vasconcelos 2008). and the efficiency of those strains to control The only difference that could explain those tetranychids on tomato greenhouses should be variations beyond the species might be the given later. photoperiod, which was 16/ 8 (L/ D) in our experiment, and 12/ 12 or 14/ 10 (L/ D) on the others (Overmeer 1985). Values of rm reported by Acknowledgements Furtado (2007) for P. longipes B, and by Badii (1984) for P. longipes SA, both reared on bean, are The PhD grant for the whole study in which this very similar to what was obtained in the present paper is included is coming half from the French study for P. longipes C feeding on T. urticae on ANRT (Association Nationale de la Recherche bean, with 0.320, 0.366 and 0.368 female/ female/ Technique) and half from Koppert BV, The day, respectively. Similar or lower figures were Netherlands. The authors would like to thank found for other species feeding on T. urticae on particularly Pr. Salvatore Ragusa for its help in bean, ranging from 0.373 to 0.193 female/ female/ providing the strain of Phytoseiulus longipes, and day, for P. persimilis and P. macropilis, respectively Alain Migeon for providing Tetranychus evansi. (Escudero & Ferragut 2005; Silva et al. 2005). However, the low rm found in the present References experiment for P. longipes C feeding on T. urticae Badii M.H., McMurtry J.A. 1983. Effect of different foods on tomato can suggest that the item T. urticae/ on development, reproduction and survival of tomato is not suitable for this strain, thus that no Phytoseiulus longipes [Acarina: Phytoseiidae]. effective control of T. urticae could be considered Entomophaga 28, 161‐166. by releasing this predaceous mite on tomato greenhouses. This is the first report of a study on the biology of a

Montpellier ‐ 21‐25 July 2008 466 Badii M.H., McMurtry. J.A. 1984. Life history of and life Ho C. C., Wang S. C., Chien Y. L. 2005. Field observation table parameters for Phytoseiulus longipes with on 2 newly recorded spider mites in Taiwan. Plant comparative studies on P. persimilis and Protection Bulletin (Taipei) 47 4, 391‐402. Typhlodromus occidentalis (Acari: Phytoseiidae). Humber R. A., Moraes G. J. de 1981. Natural infection of Acarologia XXV, 111‐123. Tetranychus evansi (Acarina: Tetranychidae) by a Badii M.H., McMurtry J.A., Flores A.E. 1999. Rates of Triplosporium sp (Zygomycetes: Entomophtorales) in development, survival and predation of immatures Northeastern Brazil. Entomophaga 26, 421‐425. stages of Phytoseiulus longipes (Acari: Koller M., Knapp M., Schausberger P. 2007. Direct and Mesostigmata: Phytoseiidae). Experimental and indirect adverse effects of tomato on the predatory Applied Acarology 23, 611‐621. mite Neoseiulus californicus feeding on the spider Birch L.C. 1948. The intrinsic rate of natural increase of mite Tetranychus evansi. Entomol. Experimental and an insect population. Journal of Animal Ecology 17, Applied Acarology 125, 297‐305. 15‐26. Mabeya J., Knapp M., Nderitu J. H., Olubayo F. 2003. Blair B.W. 1989: Laboratory screening of acaricides Comparaison de l'efficacité de Oberon against Tetranychus evansi Baker and Pritchard. Crop (Spiromefisen) avec celle d'autres acaricides dans la Protection 8, 212‐216. lutte contre les acariens rouges (Tetranychus evansi th Bolland H.R., Vala F. 2000. First record of the spider mite Baker & Pritchard) sur la tomate. 15 Biennal Tetranychus evansi (Acari: Tetranychidae) from Congress of the African Association of Insect Portugal. Entomologische Berichten 60 (9),180. Scientists (AAIS), Nairobi, Kenya, 2003. (French) Castagnoli M., Nannelli R., Simoni S. 2006. Un nuovo Maluf W. R., Campos G. A., Cardoso M. G. 2001. temibile fitofago per la fauna italiana: Tetranychus Relationships between trichome type and spider evansi (Baker e Pritchard) (Acari Tetranychidae). mite (Tetranychus evansi) repellence in tomatoes Informatore Fitopatologico 5, 50‐52. (in Italian) with respect to foliar zingiberene contents. Euphytica 121, 73‐80. Escudero L. A., Ferragut F. 2005. Life history of predatory mites Neoseiulus californicus and Phytoseiulus Maia A., Luiz A. J. B., Campanhola C. 2000. Statistical persimilis (Acari: Phytoseiidae) on four spider mite inference on associated fertility life table parameters species as prey, with special reference toTetranychus using Jakknife technique: computational aspects. evansi (Acari: Tetranychidae). Biological Control 32, Journal of Economic Entomology 93 (2), 511‐518. 378‐384. McMurtry J. A., Croft B. A. 1997. Life‐styles of phytoseiid Ferragut F., Escudero L. A. 1999. Tetranychus evansi mites and their roles in biological control. Annual Baker & Pritchard (Acari: Tetranychidae), una nueva Review of Entomology 42, 291‐321. araña roja en los cultivos horticolas españoles. Migeon A. 2005: Un nouvel acarien ravageur en France : Boletin de Sanidad Vegetal. Plagas. 25, 157‐164. (in Tetranychus evansi Baker et Pritchard. Phytoma 579, Spanish) 38‐43. (in French) Ferrero M., Moraes G.J. de, Kreiter S., Tixier M.‐S., Knapp Moraes G. J. de, Lima H. C. 1983. Biology of Euseius M. 2007. Life tables of the predatory mite concordis (Chant) (Acarina: Phytoseiidae) a predator Phytoseiulus longipes feeding on Tetranychus evansi of the tomato russet mite. Acarologia 24, 251‐255. at four temperatures (Acari: Phytoseiidae, Moraes G. J. de, McMurtry J. A. 1985a. Chemically Tetranychidae). Experimental and Applied Acarology mediated arrestment of the predaceous mite 41, 45‐53. Phytoseiulus persimilis by extracts of Tetranychus Furtado I.P., Moraes G.J. de, Kreiter S., Knapp M. 2006. evansi and Tetranychus urticae. Experimental and Search for effective natural enemies of Tetranychus Applied Acarology 1, 127‐138. evansi in south and southeast Brazil. Experimental Moraes G. J. de, McMurtry J. A. 1985b. Comparison of and Applied Acarology 40 (3‐4), 157‐174. Tetranychus evansi and T. urticae [Acari: Furtado I. P., Moraes G. J. de, Kreiter S., Tixier M.‐S., Tetranychidae] as prey for eight species of Phytoseiid Knapp M. 2007. Potential of a Brazilian population of mites. Entomophaga 30, 393‐397. the predatory mite Phytoseiulus longipes as a Moraes G. J. de, McMurtry J. A. 1986. Suitability of the biological control agent of Tetranychus evansi (Acari: spider mite Tetranychus evansi as prey for Phytoseiidae, Tetranychidae). Biological Control 42, Phytoseiulus persimilis. Entomologia Experimentalis 139‐147 et Applicata 40, 109‐115. Gonçalves L. D., Maluf W. R., Cardoso M. G., Resende J. Oliveira E. E., Oliveira C. L., Sarmento R. A., Fadini M. A. T. V. de, Castro E. M. de, Santos N. M., Nascimiento I. M., Moreira L. R. 2005. Biological aspects of the R., Faria M. V. 2006. Relationship between predator Cycloneda Sanguinea (Linnaeus, 1763) zingiberene, foliar trichomes and repellence of (Coleoptera: Coccinellidae) fed with Tetranychus tomato plant to Tetranychus evansi. Pesquisa evansi (Baker & Pritchard, 1960) (Acari: Agropecuaria Brasileira 41, 267‐273. Tetranychidae) and Macrosiphum euphorbiae Gutierrez J., Etienne J. 1986. Les Tetranychidae de l'île de (Thomas, 1878) (Homoptera: Aphididae). Bioscience la Réunion et quelques‐uns de leurs prédateurs. Journal 21, 33‐39. L'Agronomie Tropicale. 41, 84‐91. (in French)

Montpellier ‐ 21‐25 July 2008 467 Overmeer W. P. J. 1985. Diapause: 95‐102. In Spider Tsagkarakou A., Cros‐Arteil S., Navajas M. 2007. First mites – Their biology, Natural Enemies and Control, record of the invasive mite Tetranychus evansi in Vol 1B. Edited by Helle W. & Sabelis M. W., Elsevier, Greece. Phytoparasitica 35, 519‐522. Amsterdam. Vasconcelos G. J. de, Moraes G. J. de, Delalibera Jr I., R Development Core Team. 2008. R. A language and Knapp M. 2008. Life history of the predatory mite environment for statistical computing. R Foundation Phytoseiulus fragariae on Tetranychus evansi and for Statistical Computing, Vienna, Austria. Tetranychus urticae (Acari: Phytoseiidae, http://www.R‐project.org Tetranychidae) at five temperatures. Experimental Resende J. T. V., Maluf W. R., Cardoso M. G., Nelson D. L. and Applied Acarology 44, 27‐36. 2002. Inheritance of acylsugar contents in tomatoes Wekesa V. W., Knapp M., Maniania N. K., Bog, H. I. 2006. derived from an interspecific cross with the wild Effects of Beauveria bassiana and Metarhizium tomato Lycopersicon pennellii and their effect on anisopliae on mortality, fecundity and egg fertility of spider mite repellence. Genetic and Molecular Tetranychus evansi. Journal of Applied Entomology Research 1, 106‐116. 130, 155‐159. Resende J. T. V., Maluf W. R., Cardoso M. G., Faria M. V., Wekesa V. W., Maniania N. K., Knapp M., Boga H. I. Gonçalves L. D., Nascimento I. R. do 2008. Resistance 2005. Pathogenicity of Beauveria bassiana and of tomato genotypes with high level of acylsugars to Metarhizium anisopliae to the tobacco spider mite Tetranychus evansi Baker & Pritchard. Scientia Tetranychus evansi. Experimental and Applied Agricola 65, 31‐35. Acarology 36, 41‐50. Rosa A. A., Gondim Jr. M. G. C., Fiaboe K. K. M., Moraes Wekesa V. W., Moraes G. J. de, Knapp M., Delalibera Jr I. G. J. de, Knapp M. 2005. Predatory mites associated 2007. Interactions of two natural enemies of with Tetranychus evansi Baker & Pritchard (Acari: Tetranychus evansi, the fungal pathogen Neozygites Tetranychidae) on native solanaceous plants of floridana (Zygomycetes: Entomophtorales) and the coastal Pernambuco State, Brazil. Neotropical predatory mite, Phytoseiulus longipes (Acari: Entomology 34, 689‐692. Phytoseiidae). Biological Control 41, 408‐414. Sarmento R. A., Oliveira H. G. de, Holtz A. M., Silva S. M. Zhang Z. Q. 2003. Mites of greenhouses, Identification, da, Serrão J. E., Pallini A. 2004. Fat body morphology Biology and Control. CABI, London, 244 p. of Eriopis connexa (Coleoptera, Coccinelidae) in function of two alimentary sources. Brazilian Archives of Biology and Technology 47, 407‐411. Silva F. R. da, Vasconcelos G. J. N., Gondim Jr M. G. C., Oliveira J. V. 2005. Exigências térmicas e tabela de vida de fertilidade de Phytoseiulus macropilis (Banks) (Acari: Phytoseiidae). Neotropical Entomology 34 (2), 291‐296 (in Brazilian). Takahashi F., Chant D. A. 1992. Adaptive strategies in the genus Phytoseiulus Evans (Acari: Phytoseiidae): I. Developmental times. International Journal of Acarology 18, 171‐176.

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