Population dynamics of Aceodromus convolvuli (Acari:Mesostigmata: Blattisociidae) on spontaneous plantsassociated with Jatropha curcas in central Brazil

Wilton P. Cruz • Renato A. Sarmento • Marcal Pedro-Neto •

Adenir V. Teodoro • Diego M. Rodrigues • Gilberto J. de Moraes

Received: 5 December 2013 / Accepted: 5 June 2014 / Published online: 19 June 2014� Springer International Publishing Switzerland 2014

Abstract Spontaneously growing plants are commonly considered competitors of cul-

tivated plants. Owing to the lack of specificity of many arthropods, spontaneous plants may

be attacked by the same arthropods that attack cultivated plants and they may also harbor

natural enemies of organisms harmful to cultivated plants. Aceodromus convolvuli Muma

(Blattisociidae) has been reported recently in relatively large numbers in Tocantins state,

central Brazil, mostly on Helicteres guazumifolia Kunth (Malvaceae). Very little has been

reported about the population dynamics of blattisociid mites under field conditions. The

objective of this work was to study the population dynamics of A. convolvuli in Gurupi,

Tocantins state, to evaluate its possible interaction with associated mites. Monthly samples

were taken from leaves of the 11 most abundant and frequent spontaneous plants in a

Jatropha curcas L. (Euphorbiaceae) plantation. About 96.5 % of the specimens of A.

convolvuli were collected in the rainy season. The patterns of variation of the population of

A. convolvuli and of predators belonging to the family Phytoseiidae were similar, but A.

convolvuli was much more numerous than all phytoseiid specimens combined. Highly

W. P. Cruz � R. A. Sarmento (&) � M. Pedro-NetoUniversidade Federal de Tocantins (UFT), PO Box 66, Gurupi, State of Tocantins, Brazile-mail: rsarmento@uft.edu.br

W. P. Cruz � G. J. de MoraesPrograma de Pos-graduacao em Entomologia Agrıcola, Universidade Estadual Paulista Julio deMesquita Filho, UNESP, Jaboticabal, State of Sao Paulo, Brazil

A. V. TeodoroEmbrapa Tabuleiros Costeiros, Av. Beira Mar 3250, PO Box 44, Jardins, Aracaju, State of Sergipe,Brazil

D. M. RodriguesDepartamento de Ciencias Agrarias, Universidade Federal do Para (UFPA), Maraba, State of Para,Brazil

G. J. de MoraesDepartamento de Entomologia e Acarologia, Escola Superior de Agricultura ‘‘Luiz de Queiroz’’,Universidade de Sao Paulo (ESALQ/USP), Piracicaba, State of Sao Paulo, Brazil

123

Exp Appl Acarol (2014) 64:309–319DOI 10.1007/s10493-014-9828-5

significant correlations were observed between A. convolvuli densities and relative

humidity or diversity of spontaneous plants. When only mites on H. guazumifolia were

considered, highly significant correlation was also observed between densities of A. con-

volvuli and of mites of the family Tetranychidae. The results suggested that A. convolvuli

could be a predator of tenuipalpid and/or tetranychid mites. Studies about its biology are

needed to determine its preferred food sources and potential as biological control agent.

Keywords Ecology � Predation � Biological control � Blattisociid mite

Introduction

Spontaneously growing plants are often considered competitors of cultivated plants, able to

reduce the yield of the latter, being in those cases referred to as invasive plants or weeds.

Because of the lack of specificity of many arthropods, spontaneous plants may be attacked

by the same arthropods that attack cultivated plants and could thus serve as reservoirs of

pest organisms. However, they may also harbor natural enemies of organisms that are

harmful to cultivated plants (Altieri et al. 2003; Bellini et al. 2005).

The main mites of the order Mesostigmata found on plants are predators belonging to

the family Phytoseiidae. Some species of this family are considered to be important to the

control of several plant pests (McMurtry and Croft 1997; Gerson et al. 2003). Much less

common on plants are the members of the family Blattisociidae, which were previously

included in the family Ascidae (Lindquist and Evans 1965; Halliday et al. 1998; Lindquist

et al. 2009). Several blattisociids have been mentioned to prey on arthropods (Gerson et al.

2003; Lindquist et al. 2009).

Aceodromus Muma is a mite genus originally included in Phytoseiidae (Muma 1961)

and later transferred to Blattisociidae (Lindquist and Chant 1964). Aceodromus convolvuli

Muma was described from specimens collected in Florida, United States of America, on

leaves of Convolvulus sp. (Muma, 1961). It was later reported by Lindquist and Chant

(1964) in Jamaica on leaves of eggplant and Crotalaria sp., and by Fadamiro et al. (2009)

in Alabama, USA, on unidentified plants in citrus orchards. Aceodromus convolvuli was

first reported in Brazil by Moraes et al. (1993), on spontaneous plants growing in several

states of the northeastern part of the country. It was also reported on coffee in Sao Paulo

State, Brazil (Mineiro et al. 2006).

More recently, A. convolvuli was reported from Tocantins state, in central Brazil, in a

study conducted by Cruz et al. (2012) on the mite fauna of physic nut Jatropha curcas L.

(Euphorbiaceae) and associated spontaneous plants. Jatropha curcas has been considered

for extended cultivation in several tropical countries, mainly for biodiesel production

(Openshaw, 2000).

In the work conducted by Cruz et al. (2012), 1,141 specimens of A. convolvuli were

collected on samples of 11 species of spontaneous plants. About 69 % of those specimens

were found on Helicteres guazumifolia Kunth (Malvaceae). This plant has been reported

from southern Mexico to the central region of Brazil, in open, secondary and semidecid-

uous forests, gallery forests, pastures, zones of periodic fires and clearings, dry or moist

thickets and grassy or bushy slopes (Goldberg 2009). In Venezuela, this plant has been

reported mainly in open habitats with dry, sandy and rocky soils (Rondon and Cumana-

Campos 2007).

310 Exp Appl Acarol (2014) 64:309–319

123

The objective of this work was to conduct an analysis of the population dynamics of A.

convolvuli, based on the samplings conducted by Cruz et al. (2012), evaluating its possible

interaction with the associated mites on those 11 spontaneous plant species. This work is

one of the rare efforts to evaluate the population dynamics of species of the ascid sense

Lindquist and Evans (1965). Apparently, the only other study of this type refers to the

blattisociid Lasioseius parberlesei Bhattacharyya in Taiwan (Tseng 1984), where it was

considered an effective predator of Steneotarsonemus spinki Smiley (Tarsonemidae) on

rice. Thus, not much can be anticipated in terms of the response of A. convolvuli to the

varying climatic factors, or in terms of its possible natural prey. However, as other blat-

tisociid mites have a preference for humid environments, we anticipated that the same

occurs with A. convolvuli.

Materials and methods

The data used for the analysis presented here were collected in the study conducted by

Cruz et al. (2012). The work involved the evaluation of A. convolvuli and associated mite

species on spontaneous plants from a 4-year old cultivation of J. curcas of about 0.5 ha, in

Gurupi (11�4802900S, 48�5603900W; 280 m altitude), Tocantins state, in a subtropical region

with well defined rainy (November–April) and dry (May–October) seasons. The natural

vegetation of this locality is classified as ‘‘Cerrado’’, a very diverse savannah type of biome

covering slightly over 20 % of the Brazilian territory, where annual rainfall ranges between

800 and 2,000 mm, but with a pronounced dry season (Ratter et al. 1997). The experi-

mental field was mainly surrounded by pasture (mostly Brachiaria sp., Poaceae), except

for one side, edging an area of natural vegetation.

The height of the spontaneous plants varied from about 15 cm (dry season) to about

120 cm (wet season). They were not controlled during the conduction of the study or in the

previous year. No fertilization or pesticide spray was done during the study or in the period

of 2.5 years before it started (Cruz et al. 2012).

Mite sampling

Samples were taken from 15 sites (9 m2 each) randomly delimited within the experimental

field. Eleven plant species were selected for the evaluations, based on their abundance and

frequency in the area (Cruz et al. 2012). The selection was done by counting the plants of

each species within two quadrats (0.25 m2) randomly thrown in each sampling site (Mu-

eller-Dombois and Ellenberg 1974; Cruz et al. 2012). Selected species were those with

abundance of at least 2.5 plants per quadrat and which were present with a frequency of at

least 10 % of the quadrats. The same selected plant species were sampled throughout the

work, regardless of the variation in their abundance or frequency along the time. Identi-

fication of spontaneous plants was conducted with the use of a manual (Lorenzi 2006) and

the help of M. Ignacio, a botanist. The selected plants were: Fabaceae—Calopogonium

mucunoides Desv.; Lamiaceae—Hyptis suaveolens (L.) Poit.; Malvaceae—Peltaea riedelii

(Gurke) Standl., Sida urens L., Sida cordifolia L., Sida rhombifolia L., Urena lobata L.,

H. guazumifolia, Waltheria americana L.; Poaceae—Andropogon gayanus Kunth and

Urochloa mutica (Forssk.) T.Q. Nguyen (Angiosperm Phylogeny Group 2009). The

diversity of spontaneous plants was estimated at each sampling date by calculating the

Shannon–Wiener (H0) index with DIVES software version 2.0. Sampling of spontaneous

plants for diversity estimation was conducted using the methodology proposed for Mueller-

Exp Appl Acarol (2014) 64:309–319 311

123

Dombois and Ellenberg (1974), described above. Subsequently, the spontaneous plants’

richness and abundance were obtained.

The samples were collected once between the 20th and 25th of each month, from

February 2010 to January 2011. At each sampling date, 10 (Fabaceae, Lamiaceae and

Malvaceae) or 20 (Poaceae) leaves of each selected plant species present in each sampling

site were collected, constituting a sample. Thus, for each species, the number of samples

taken at each sampling date varied according to its respective frequency at that sampling

date, for a minimum of two and a maximum of 11 samples. The leaves of each species

from each sampling site were placed in a plastic bag that in turn was placed in a cool box

(about 20 �C) and transferred to the laboratory, where they were maintained in a refrig-

erator (about 10 �C) for up to 5 days, until examined under a stereomicroscope. For

quantification and identification, mites were mounted in Hoyer’s medium; however, when

a certain species was too abundant, the mites were counted and only part of the specimens

was mounted for confirmation of the identification. Identification of mite species was

carried out by the third and the last authors, using the original descriptions and relevant

complementary descriptions of the species of each group.

Leaf area was estimated at each sampling date. Climatic data (daily temperature, rel-

ative humidity and rainfall) were obtained from ‘‘Estacao Meteorologica de Gurupi (IN-

MET)’’, at Universidade Federal do Tocantins campus, about 20 km from the experimental

site. The analyses were conducted by taking into account monthly average temperature and

relative humidity and monthly total rainfall, in all cases for the period preceding each

sampling date (i.e., the most recent monthly reading was taken).

Statistical analysis

To estimate the main factors related to the fluctuation of A. convolvuli throughout the work,

data distribution was initially checked with Shapiro–Wilk normality test, using the Shap-

iro.test function procedure of R program. Since many of the variables did not have normal

distribution, Spearman’s test was used to determine simple correlations between variables

(population densities of A. convolvuli, phytoseiids, tetranychids and tenuipalpids, as well as

indexes of plant diversity, rainfall, relative humidity and temperature). This was done with

the use of the Spearman’s rank correlation (rho) function procedure or R program. A

multiple regression analysis relating the population densities of A. convolvuli with the

variables showing significant simple correlation with densities of A. convolvuli and not

related among themselves was performed, using ‘‘model’’ of R to obtain the coefficients of

the function and Summary (model) to determine the level of significance of the variables and

of the model as a whole, and R2 of the model (R Development Core Team 2006).

Results

A re-examination of the mites collected showed the need to correct or complement the

identification reported in Cruz et al. (2012), as indicated in Table 1.

Relation between Aceodromus convolvuli and environmental factors in rainy and dry

seasons

Considering all plants together, about 96.5 % of the specimens of A. convolvuli were

collected in the rainy season (Table 2). In the dry season, that species was only found on H.

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guazumifolia and C. mucunoides, at very low numbers. Total rainfall, average temperature

and average relative humidity were 964 mm, 26 ± 0.6 �C and 80 ± 1.1 % in the wet

season, and 80 mm, 26 ± 1.6 �C and 59 ± 3.9 % in the dry season, respectively.

Dynamics of Aceodromus convolvuli on all spontaneous plants

When all spontaneous plant species were considered together, the densities of A. convolvuli

increased between February and April, when the maximum level was reached (Fig. 1a).

Densities reduced drastically to nearly zero in June, remaining so until September; it again

increased progressively but slightly until November, remaining about the same level until

next January. The pattern of variation of the phytoseiid population was similar, except that

the decline of the population after reaching a peak density in April was more gradual.

Among other mite groups, the highest densities were attained by the Tydeoidea

(including Tydeidae and Iolinidae), which peaked in March, followed by the Tetranychidae

and the Tenuipalpidae, both reaching the maximum level in June (Fig. 1b). The highest

densities of A. convolvuli and phytoseiids coincided with the highest levels of plant

diversity and with the end of the rainy season (Fig. 1c).

Highly significant correlations were observed between the densities of A. convolvuli and

phytoseiids, as well as between the former and plant diversity indexes (Table 3); signifi-

cant correlations were observed between A. convolvuli densities and relative humidity. No

significant correlation between A. convolvuli densities and any of the remaining inde-

pendent variables was observed.

Owing to the significant correlations between phytoseiid densities and plant diversity as

well as between levels of relative humidity and plant diversity (Table 3), the latter

parameter was not included in the multiple regression analysis. In that analysis, the only

significant factor of the generated equation corresponded to phytoseiid densities, and the

coefficient of determination (R2) of the equation was 0.78.

Dynamics of Aceodromus convolvuli on Helicteres guazumifolia

The patterns of variation of the densities of A. convolvuli and phytoseiids were similar to

those when all spontaneous plant species were considered together (Fig. 2a). However, the

peak population density of A. convolvuli was much higher, while that of the phytoseiids

was only slightly higher. Differently from what was observed when all plants were con-

sidered together, densities of Tydeoidea were low during the whole observation period

Table 1 Correction and complementation of the identification of the mite species reported by Cruz et al.(2012), based on a re-examination of the specimens

Reported by Cruz et al. (2012) Corrected identification

Tetranychidae

Mononychellus sp.Panonychus sp.

Mononychellus planki (McGregor)Neotetranychus granifer Feres & Flechtmann

Phytoseiidae

Neoseiulus aff. mumaiNeoseiulus fallacis

Neoseiulus benjamini (Schicha)Neoseiulus pluridentatus Lofego & Moraes

Phytoseiulus sp.Typhlodromalus clavicus

Phytoseius guianensis De LeonAmblydromalus sp.

Exp Appl Acarol (2014) 64:309–319 313

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(Fig. 2b). Among the remaining mite groups, the highest levels were attained by the

tetranychids and the tenuipalpids, also in June, being these about twice as high as deter-

mined for all plants together. Yet, variations in the populations of mites other than A.

convolvuli and phytoseiids were much more erratic than observed when all plants were

considered together.

Significant correlations (Table 3) were observed between A. convolvuli densities and

phytoseiid densities, tetranychid densities, relative humidity, and plant diversity. No sig-

nificant correlation was observed between the remaining independent variables with A.

convolvuli densities.

Because of the significant correlations between phytoseiid and tetranychid densities,

phytoseiid densities and plant diversity, relative humidity and plant diversity, and tetr-

anychid densities and plant diversity (Table 3), the multiple regression analysis included

only phytoseiid and tenuipalpid densities, temperature and rainfall. Similarly to what was

determined when all spontaneous plant species were considered together, the only sig-

nificant factor of the generated equation was phytoseiid densities, and the coefficient of

determination of the equation was 0.58.

Discussion

The analyses of the numbers of A. convolvuli in each of the two distinct seasons prevailing

in Gurupi as well as of the dynamics of each variable along the evaluation period suggested

a strong influence of rainfall and relative humidity on the densities of A. convolvuli. The

significant correlation between the population level of A. convolvuli and the levels of

relative humidity confirmed that assumption, but the non-significant correlation between

the first parameter and rainfall seemed not to support it. This apparent contradiction could

be due to the non-linear relation between A. convolvuli population densities and rainfall; in

this sense, there might be a minimum rainfall threshold level below which A. convolvuli is

about uniformly and negatively affected, independent of the amount of rain. Hence,

absence or incidence of very low rainfall between May and September resulted in very low

Table 2 Seasonal occurrence of Aceodromus convolvuli on spontaneous plants, in the dry and rainy sea-sons, in Gurupi, Tocantins state, Brazil, from February 2010 to January 2011

Spontaneous plants Dry season (May–October) Rainfall season (November–April) Total

Helicteres guazumifolia 38 746 784

Calopogonium mucunoides 2 135 137

Sida urens 0 48 48

Ureda lobata 0 41 41

Waltheria americana 0 41 41

Andropogon gayanus 0 23 23

Hyptis suaveolens 0 19 19

Sida cordifolia 0 18 18

Urochloa mutica 0 16 16

Peltaea riedelii 0 8 8

Sida rhombifolia 0 6 6

Total 40 1,101 1,141

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levels or total absence of A. convolvuli in the study area. In addition, the sudden occurrence

of high rainfall after the dry season was not followed by a corresponding proportional

increase in A. convolvuli population levels, possibly because of a natural time lag required

for the food sources (possibly prey organisms) of A. convolvuli to increase with increasing

rainfall, which would then allow the subsequent increase of the population of the latter.

Fig. 1 Population densities of Aceodromus convolvuli and associated mite species on all sampled plants aswell as monthly mean levels of plant diversity and abiotic parameters, in Gurupi, Tocantins state, betweenFebruary 2010 and January 2011

Exp Appl Acarol (2014) 64:309–319 315

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With the onset of the dry season, the sharper decline of A. convolvuli population in

comparison with the phytoseiid population suggest the former to be more sensitive to lower

humidity than the latter. This pattern would be expected, as members of Ascidae sensu lato

Table 3 Significant correlations between the population densities of Aceodromus convolvuli and biotic andabiotic factors as well as between other combinations of factors for all spontaneous plants and on Helicteresguazumifolia alone, in Gurupi, Tocantins State, Brazil, from February 2010 to January 2011

Factors Spontaneous plants Helicteres guazumifolia

r P r P

Phytoseiidae 0.93 0.000009 0.80 0.002

Plant diversity 0.92 0.00001 0.87 0.0002

Relative humidity 0.66 0.01 0.71 0.01

Tetranychidae 0.66 0.01

Phytoseiidae 9 plant diversity 0.87 0.003 0.83 0.0007

Phytoseiidae 9 Tetranychidae 0.78 0.03

Relative humidity 9 plant diversity 0.60 0.04 0.61 0.04

Tetranychidae 9 plant diversity 0.61 0.03

Fig. 2 Population densities of Aceodromus convolvuli and associated mite species on Helicteresguazumifolia, in Gurupi, Tocantins state, between February 2010 and January 2011

316 Exp Appl Acarol (2014) 64:309–319

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Lindquist and Evans (1965) are usually found on aerial plant parts only in regions of

prevailing high levels of relative humidity. In a study conducted in northeastern Brazil,

Rodrigues et al. (1996) determined the significant level of association A. convolvuli with

several other mites and plants. They determined the association to be negative with the

phytoseiid Neoseiulus (=Amblyseius) idaeus Denmark & Muma and positive with the

phytoseiid Amblydromalus (=Typhlodromalus) manihoti (Moraes), species that predomi-

nate respectively under conditions of low and high humidity levels (Moraes et al. 1990). As

commonly observed in areas of the Cerrado vegetation, the region where the study was

conducted has marked alternations of very dry and very humid seasons. Thus, the prev-

alence of A. convolvuli during part of the year in Gurupi leads to the conclusion that it has

an efficient mechanism to allow survival in the dry season, though at very low population

levels (Cruz et al. 2013).

Faunistic surveys on leaves of cultivated plants or plants of the natural vegetation have

usually shown outstandingly higher abundance of phytoseiids in comparison with other

mesostigmatid mites. Thus, the finding of a larger population density of A. convolvuli in

comparison with that of all phytoseiid species combined is uncommon, deserving detailed

analysis. A question to be answered is why A. convolvuli is so abundant in the region where

the study was conducted. Among other reasons, the relatively high abundance could be

related to the presence of H. guazumifolia, onto which A. convolvuli was by far the most

common Mesostigmata. However, in which way the plant affects the mite could not be

determined, because of the absence of information about its biology. The higher proportion

of A. convolvuli than of phytoseiids was not related to a conceivable more intense sur-

veying effort on this plant, as at each sampling date the examined leaf area of that plant

species corresponded to only about 20 % of all examined leaves.

Although not rare, H. guazumifolia was not one of the most common plants considered

in this study. It was only the eighth on the list of the most abundant, with a patchy

distribution in the study area. This species is a shrub whose leaves are covered by star-

shaped trichomes, under which A. convolvuli were found. Those structures seem to offer

some protection to the predators, as suggested by the fact that it was rather difficult to

collect them with a brush during the evaluations. Several factors have been related to the

presence of trichomes, including the maintenance of lower temperature on the leaf surface,

protection from the effect of UV light and the retention of a thicker boundary layer of

higher relative humidity over the leaf surface (Wagner et al. 2004; Coder and Warnell

2013). These characteristics might favor A. convolvuli.

As reported by Ordaz et al. (2011), several studies have been conducted on the mor-

phology, phenology, ecology and biochemical characteristics of H. guazumifolia, a plant of

medicinal importance in South America. Characteristics about patterns of nectar produc-

tion, morphology of floral nectaries, nectar composition and hummingbird association were

reported by Goldberg (2009). The chemical composition of essential oils of leaves of H.

guazumifolia was studied by Ordaz et al. (2011). The authors reported that the predominant

compounds were non-terpenoid volatile secondary metabolites. The role of alternative food

such as nectar and secondary compounds to the predominance of A. convolvuli on H.

guazumifolia in this study has not been determined. To our knowledge this is the first study

of the mite fauna of this plant.

Considering the possibility of using A. convolvuli for the control of pest organisms,

future studies should include detailed evaluations on its accepted food sources. The results

of the present study suggest that items to be investigated should include tetranychid and

tenuipalpid mites. The relatively low correlation between the densities of A. convolvuli and

tetranychids as well as the non significant correlation between the densities of A.

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convolvuli and tenuipalpids might not necessarily indicate that they do not interact. Instead,

that could be a function of a complex predator–prey interactions, with non-synchronous

population fluctuations, and/or due to the concurrent confusing effect of other environ-

mental factors. The inverse patterns of variation of A. convolvuli and tenuipalpids

graphically distinguishable between February and June further supports this possibility,

with an increase in the population level of the former being followed by a (slower)

reduction of the population of the latter during the end of the rainy season, and an inverse

trend at the onset of the dry season, which, as previously discussed, seems to disfavor A.

convolvuli. Detailed studies of predator–prey interactions should be stimulated, especially

in areas where mites have been barely investigated, as in central Brazil. These may lead to

the discovery of new prospective biological control agents of this relatively little known

group of mesostigmatid mites. Considering that another blattisociid mite, Lasioseius

lindquisti Nasr & Abou-Awad, has been reported to develop and reproduce when offered

pollen of Phoenix dactylifera as food source (Momen et al. 2011), pollen of H. guaz-

umifolia should also be investigated as a possible food source for this mite.

Acknowledgments The authors thank the National Council for Scientific and Technological Development(CNPq) for financial support (Projects 620028/2008-4 and 475408/2008-0) and Coordination of Improve-ment of Higher Education Personnel CAPES-Brazil for providing scholarships to the first and third authors(Projects: PROCAD-NF and CAPES-PNPD). We thank M. Ignacio (Universidade Federal do Tocantins,Brazil) for help with the identification of part of the plants collected in this study.

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