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Ability of Phytoseiulus longipes to control spider mite
pests on tomato in European greenhouses.
Maxime Ferrero, Serge Kreiter, Marie-Stéphane Tixier
To cite this version:
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�
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, SolanaceaeIntroduction
Although many predatory mite species have been studied to control tetranychids, those pests still cause serious damages to several crops, and especially to tomato in greenhouses. In Southern Europe, the main problem has been Tetranychus
urticae Koch for many years (Zhang 2003). But
since the early nineties, an invasive pest,
Tetranychus evansi Baker & Pritchard, which seems
to be originated from South America (Gutierrez & Etienne 1986) spreads through Southern Europe (Ferragut & Escudero 1999; Bolland & Vala 2000,
Migeon 2005, Castagnoli 2006, Tsagkarakou 2007) and causes severe injuries, especially to Spanish tomato crops (Ferragut pers. comm.).
In order to get rid of T. evansi, many methods have been experimented, like pesticides (Blair 1989, Mabeya et al. 2003), plant resistance development (Maluf et al. 2001, Resende et al. 2002, Gonçalves
et al. 2006, Resende et al. 2008),
entomopathogenic fungi (Humber & Moraes 1981, Wekesa et al. 2005, 2006, 2007) and several predators (Sarmento et al. 2004, Ho et al. 2005, Oliveira et al. 2005) including predatory mites
Montpellier ‐ 21‐25 July 2008 462
(Moraes & Lima 1983; Moraes & McMurtry 1985a; Moraes & McMurtry 1985b; Moraes et McMurtry 1986; Escudero et Ferragut 2005; Rosa et al. 2005, Ferrero et al. 2007, Furtado et al. 2007, Koller et al. 2007). The only enthusiastic perspective up to now has been pointed out by Furtado et al. (2006) and Ferrero et al. (2007), with the discover and the study of a Brazilian strain of Phytoseiulus longipes Evans (named P. longipes B thereafter). Following those studies, this type I predator (McMurtry & Croft 1997), feeding preferentially on mites of the subfamily Tetranychinae, seems actually to be specific of T. evansi (Furtado et al. 2007). More recently, a Chilean strain of this species (written P.
longipes C thereafter) has been found (Ragusa
pers. comm.). In order to use these strains in biological control for controlling T. evansi, further biological studies are required. This paper aims to determine the life history of P. longipes C on several plant substrates/ prey conditions.
Material and Methods
Species and strains studied
The strain of P. longipes studied was obtained from a colony initiated with specimens collected in Chile, fed with T. urticae, in Nogal, Los Andes, Valparaiso Region (Ragusa, pers. comm.). Mites were fed with all stages of T.
urticae, offered on leaves of Phaseolus vulgaris L. cv. Contender, placed underside up
in rearing units constituted of plastic trays (10 × 15 cm) bordered with water‐saturated cotton to avoid mite escapes and to maintain the turgescence of the leaves. Rearing units were placed in climatic units at 25 ± 2 °C, 75 ± 10 % RH and 16/ 8 [L/ D]. The T. evansi stock colony was initiated with specimens collected from a tomato screenhouse at Saint‐Jeannet (Alpes‐Maritimes, 06, France) in October 2007 (Migeon pers. comm.), and reared on
Lycopersicon esculentum Miller in rearing
units similar to those described above.
The T. urticae stock colony initiated with specimens collected in Montpellier, France, was reared on P. vulgaris cv. Contender in a greenhouse.
Immature development
Experimentations were performed at 25 ± 2 °C, 75 ± 10 % RH and 16/ 8 [L/ D]. Three items were tested, each one being characterised by a prey (T.
evansi or T. urticae) and a plant substrate (tomato
or bean): T. evansi / tomato, T. urticae / bean and
T. urticae / tomato. The combination T. evansi /
bean has not been tested because this plant was unsuitable for rearing our strain of T. evansi. Groups of five prey females were placed using a thin paintbrush in experimental units, consisting of a leaf disk of a plant substrate (2 cm in diameter) placed underside up onto a moist disk of filter paper, inside a Petri dish (2 cm in diameter, 1 cm high). One day later, eggs of P. longipes (between 0 and 6 hours old) obtained from the intermediate stock colony (consisting in 30 to 50 females reared on a leaf of P. vulgaris cv. Contender placed in a plastic tray as described above) were transferred to each experimental unit (one egg/ unit); then closed with a transparent plastic film. To maintain humidity, distilled water was added on the filter paper every day. Periodically (once a week on tomato, twice on bean) P. longipes individuals were transferred to new leaves infested with preys as previously reported.
Observations were carried out every 12 hours to determine the duration and the survivorship for each stage.
Reproduction
Recently emerged adult P. longipes females obtained were transferred to new experimental units. A male taken from the stock colony was then added to each unit containing one female, and a new one was added for every male that died or escaped. At least 30 couples were observed. Daily observations were conducted to determine female fecundity and survivorship. The eggs laid were placed daily in a single unit and reared to adulthood to determine the secondary sex ratio (female percentage of the studied female cohort offspring).
Life Table
The life table was constructed considering the females of the cohort studied. The net reproductive rate (Ro), the mean generation time
(T), the intrinsic rate of increase (rm), the doubling
generation time (Dt), and the finite rate of increase
(λ) were calculated using the method
recommended by Birch (1948): Ro = Σ (lx × mx) T = Σ (x × lx × mx)/ Σ (lx × mx) rm = Ln (Ro) / T Dt = Ln (2) / rm λ = exp (rm)
day when eggs had been laid), lx, the cumulative
female survivorship, and mx, the number of female
descendants per female at x.
Calculation of a corrected rm value was performed
by iteration. The method, aiming to find rm for
which (1 ‐ Σ exp (‐rm × x) × lx × mx) is minimal, was
given by Maia et al. (2000).
Analysis of variance (ANOVA) and related Tukey HSD mean comparison tests were performed to determine differences between duration of the immature phases and adult stages between the different items tested.
The rm iteration and statistical analysis were
computed with R (R project 2008).
Results
Immature development
Egg to adult duration ranged from 4.10 to 4.21 days, for the T. evansi/ tomato and T. urticae/ tomato conditions, respectively (Table 1). The egg stage was the longest, varying from 1.80 to 1.96 days. The larval stage was the shortest, varying from 0.35 to 0.55 days. No statistical analysis could be performed for the item T. evansi/ tomato for the protonymphal, deutonymphal and egg to adult stages because too many individuals had died prematurely (11.1 % of the immatures tested
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).
Item Stage
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
N 27 41 44 Immature survival (%) 11.1 99.8 90.9 Ft > F (2, 80) = 2.91; P= 6.03 × 10‐2 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). Plant substrate Stage Bean Tomato F (df1, df2) (α = 0.05)
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
‐1
Post‐oviposition 0.30 (0.46) a 1.21 (1.64) b Ft < F (1, 40) = 6.10; P = 1.79 × 10‐2
Longevity 8.95 (2.18) a 8.98 (2.62) a
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 (1, 51) = 6.00 × 10‐4; P = 9.81 × 10‐1 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 464 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). Demographic parameter Substrate
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
survived). No significant differences were found between the two other conditions tested for the egg to adult and deutonymph durations. For the three other immature stages (i.e. egg, larva and protonymph), significant differences were always found between the items T. urticae/ bean and T. urticae/ tomato.
Reproduction
Adult phase durations and oviposition rates were observed only when P. longipes was feeding on T.
urticae, on bean or tomato (Table 2), because of
the high mortality of P. longipes when fed with T.
evansi. The longevity was not significantly different
between the two conditions, varying from 8.95 and 8.98 days, on bean and tomato, respectively. The pre‐oviposition period ranged from 1.22 to 0.95 days, oviposition from 3.74 to 2.63 days and post‐ oviposition from 0.30 to 1.21 days, on bean and
tomato, respectively. The only significant
difference found between these two conditions was for the post‐oviposition period. The mean oviposition rate and total oviposition were highest when P. longipes was fed with T. urticae on bean than on tomato, being 1.34 eggs/ female/ day and 6.77 eggs/ female, respectively.
Life tables
Calculated life table parameters are given in Table 3. Congruently with the previous observations, the best features were obtained for the item T.
urticae/ bean. The intrinsic rate of increase
(iterated) was more than three times higher on bean than on tomato, while the doubling time and the net reproductive rate were almost three times lower on bean than on tomato. The calculated mean generation time was 4.04 and 2.91 days, and the finite rate of increase 1.40 and 1.12, on bean and tomato, respectively.
Discussion, conclusions and perspectives
This study has been performed in order to determine first the ability of the predaceous mite
P. longipes C to develop and reproduce fed with
tetranychids on tomato, through the analysis of its life parameters The second goal was to compare the biological characteristics of the studied strains of this species. Up to 2007, experiments had been performed on two other strains of this predator: a South‐African strain (reported as P. longipes SA thereafter) and a Brazilian strain (Badii & McMurtry 1983, 1984; Moraes & McMurtry 1985b; Takahashi & Chant 1992, Badii et al. 1999). It has been reported that the South‐African strain is unable to control T. evansi in the conditions tested, even if promising results had been found to control
Tetranychus pacificus (McGregor) (Badii et al.,
1999). On the opposite, the results obtained for the Brazilian strain, discovered in 2005 (Furtado et al. 2006), are enthusiastic for the control of T.
evansi but also of T. urticae on tomatoes (Ferrero et al. 2007; Furtado et al. 2007.
Concerning the developmental phase of P. longipes C, several points could be pointed out. At first, for the item T. evansi/ tomato, only 3 individuals out of 27 reached the adult stage, and none of these 3 mites survived enough to mate. These conditions thus seem to be unsuitable for the Chilean strain of
P. longipes, which is a key difference to notice with P. longipes B, which seems specific to T. evansi
(Furtado et al. 2007). For the other conditions tested in the present paper (see table IV), egg to adult durations were lower to what has been observed previously with 4.7 and 4.8 days while P.
longipes B was feeding on T. evansi and T. urticae,
respectively, on Canavalia ensiformis (L.) (Furtado
et al. 2007), 4.9 days while P. longipes B was
feeding on T. evansi on Solanum americanum Miller (Ferrero et al. 2007) and 5.23 days for P. longipes SA feeding on T. pacificus on bean (Badii & McMurtry 1984). The Chilean strain of P. longipes thus seems to develop faster than the other strains feeding on T. urticae.
The whole immature phase durations are of the same order of magnitude or longer for the other
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. Refer ence Fer rer o et al . 2 007 Furtado et al . 2 007 Badi i & McMurtry 19 84 Silva et al . 2005 Escude ro & Fe rr agu t 20 05 Va scon cel o s et al . 20 08 Intr insic rat e of inc rease rm (f e m al e/ female/ day ) 0. 29 3 0. 32 0. 36 3 0. 36 6 0. 19 3 0. 11 6 0. 37 3 0. 27 3 Longevity in days 20. 3 29. 7 31. 1 34. 1 2 44 NA NA 23. 3 Immat u re development dura tion in days 4. 9 4. 8 4. 7 5. 23 4. 8 6. 91 4. 16 5. 5 Clim atic condition s 25 ± 2 °C, 80 ± 10 % RH, 12/ 12 L/ D 25 ± 1 °C, 83 ± 12 % RH, 12/ 12 L/ D 25 ± 1 °C, 80 ± 7 % RH, 14 / 10 L/ D 26 °C , 60 ± 5 % RH , 12 / 12 L/ D 25 ± 1 °C, 70 ‐80 % RH, 12 / 12 L/ D 25 ± 0. 8, 12 / 12 L/ D Plant substrate Solan u m a m eric an um C ana val ia en siformis P. lunat us C. en siformis P. vul gar is S. ame ri can u m Prey sp ecies Te tr any chus ev ansi T. ur ti cae T. e vansi T. paci fi cus T. ur ti cae T. e vansi T. ur ti cae T. ur ti cae P h yt ose iul us species P. lo ngi pe s B P. lo ngi pe s SA P. m a cr op ilis P. p ersim ilis P. f ragar ia e
Montpellier ‐ 21‐25 July 2008 466
fragariae Denmark & Schicha feeding on T. urticae
on S. americanum (Vasconcelos et al. 2008), 4.8 days for Phytoseiulus macropilis (Banks) feeding on
T. urticae on C. ensiformis (Silva et al. 2005) and
6.91 and 4.16 days for Phytoseiulus persimilis Athias‐Henriot feeding on T. evansi and T. urticae
on bean (Escudero & Ferragut 2005). Results
obtained here on the developmental phase of P.
longipes C showed that this strain is not able to
develop, thus to control, T. evansi on tomato, but can develop in the same way on tomato and bean while feeding on T. urticae. At last, as defined by McMurtry & Croft (1997), type I phytoseiids, belonging to the genus Phytoseiulus, do not need food until they reach the protonymphal stage. This has also been observed during the present experiment for the Chilean strain of P. longipes Adult phase durations of the Chilean strain of P.
longipes were calculated only with T. urticae as
prey. These durations were very short compared to all the data previously compiled (table IV). Longevity was only 8.95 and 8.98 days in our data for items T. urticae/ bean and T. urticae/ tomato, respectively, while Ferrero et al. (2007) showed a 20.3 days longevity for P. longipes B and Badii (1984) a 34.12 days longevity for P. longipes SA. For the other Phytoseiulus species studied feeding on T. urticae at the same temperatures than in the
present study, longevity is also variable, from 44.0
and 23.3 days, for P. macropilis and P. fragariae, respectively (Silva et al. 2005; Vasconcelos 2008). The only difference that could explain those variations beyond the species might be the 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
Furtado (2007) for P. longipes B, and by Badii (1984) for P. longipes SA, both reared on bean, are very similar to what was obtained in the present study for P. longipes C feeding on T. urticae on bean, with 0.320, 0.366 and 0.368 female/ female/ day, respectively. Similar or lower figures were found for other species feeding on T. urticae on bean, ranging from 0.373 to 0.193 female/ female/ day, for P. persimilis and P. macropilis, respectively (Escudero & Ferragut 2005; Silva et al. 2005).
However, the low rm found in the present
experiment for P. longipes C feeding on T. urticae on tomato can suggest that the item T. urticae/ tomato is not suitable for this strain, thus that no effective control of T. urticae could be considered by releasing this predaceous mite on tomato greenhouses.
This is the first report of a study on the biology of a
Chilean strain of P. longipes. This work showed results different than those already obtained for other strains of this species. Phytoseiulus longipes C was, as the SA strain, unable to complete its development feeding on T. evansi on tomato, while the B strain could. Those observations lead to the first major conclusion of this work which is that P. longipes C would be unable to control T.
evansi populations in tomato greenhouses. Great
differences in longevity had been also pointed out between the Chilean strain and the two others. These differences could be due to differences in the setup (photoperiod), or to the strain itself.
Values of rm showed that P. longipes C would be
able to develop and reproduce well feeding on T.
urticae on bean, but apparently not enough
feeding on T. urticae on tomato to settle an enthusiastic prognosis for its use in biological control.
In this paper, particular attention was paid to the differences between the three strains of P.
longipes already studied: differences in the
suitability of plant substrate and in prey items, in differences in longevity and oviposition at the same conditions. All these observations lead the authors to settle a whole study to compare those strains in combined conditions. Along with more biological studies, other tools like genetic markers or morphological analysis are presently being used to determine and explain some of those differences. Many conclusions on the usefulness and the efficiency of those strains to control tetranychids on tomato greenhouses should be given later.
Acknowledgements
The PhD grant for the whole study in which this paper is included is coming half from the French ANRT (Association Nationale de la Recherche Technique) and half from Koppert BV, The Netherlands. The authors would like to thank particularly Pr. Salvatore Ragusa for its help in providing the strain of Phytoseiulus longipes, and Alain Migeon for providing Tetranychus evansi.
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