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Diel Activity Patterns and Microspatial Distribution of the Harvestman Phalangium Opilio (Opiliones, Phalangiidae) in Soybeans

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Diel Activity Patterns and Microspatial Distribution of the Harvestman Phalangium Opilio (Opiliones, Phalangiidae) in Soybeans 2005. The Journal of Arachnology 33:745±752 DIEL ACTIVITY PATTERNS AND MICROSPATIAL DISTRIBUTION OF THE HARVESTMAN PHALANGIUM OPILIO (OPILIONES, PHALANGIIDAE) IN SOYBEANS Cora M. Allard and Kenneth V. Yeargan: Department of Entomology, University of Kentucky, S-225 Ag Science North, Lexington, KY 40506 USA. E-mail: [email protected] ABSTRACT. Phalangium opilio L. is a polyphagous predator frequently found in agricultural habitats. Although the potential importance of P. opilio's feeding on pests has been recognized, little is known about its activity patterns or its within-plant distribution in crops. We determined diel activity patterns and microspatial distribution in small, fenced arenas in soybean ®elds. The fenced arenas allowed us to track known numbers of particular size categories of P. opilio for each 24 h trial. Phalangium opilio were separated into the following categories based on body size and sex: medium-sized nymphs, large-sized nymphs, adult females and adult males. Medium-sized nymphs occupy the bottom and middle portions of plants regardless of time of day; they remain still during the day, but they exhibit leg palpating behavior from 21:00±01:00 h. Large-sized nymphs rest in the bottom and middle portions of plants during the day, but they walk and palpate on the ground from 21:00±01:00 h. Adult females rest in the bottom, middle and top portions of plants during the day, and they walk and palpate on the ground from 21:00±01:00 h. Adult males remain stationary in the bottom, middle and top portions of plants during the day, but they walk on the ground from 21:00±04:00 h. Keywords: Predator, behavior, microhabitat separation The microspatial distribution and diel activ- among habitats. When sparse shrub cover is ity patterns of predators in crops affect the present, 88% of P. opilio are found in shrubs prey they encounter, which potentially affects and brushy vegetation, but with dense cover, their value in biological control. When differ- the highest percentage of P. opilio is found on ent instars of the same species separate their the ground layer (Edgar 1980). Cloudsley- location and activities spatially and temporal- Thompson (1968) also observed P. opilio to ly, the separation may reduce cannibalism primarily inhabit low vegetation or grass and and/or intraspeci®c competition for resources. other herbaceous plants, but he stated that the All these factors have the potential to affect early instars only occur on the ground. Several the population dynamics of arthropod species hypotheses have been presented to explain mi- in agricultural systems. Predatory members of crohabitat separation in Opiliones. Opilionids the group Opiliones are sometimes overlooked may be found on vegetation to eliminate com- in crops. One such predator is Phalangium op- petition with strict ground predators (Halaj & ilio L. 1758. We studied P. opilio's micro- Cady 2000). It also has been hypothesized that spatial distribution and diel activity because of the vertical expansion of the distribution of its potential importance in biological control late instars is due to the need for larger prey, of soybean pests (Anderson 1996; Pfannen- more moving space, mating, and/or different stiel & Yeargan 2002). temperature and humidity requirements (San- Microhabitat separation of different life key 1949; Todd 1949). Not all individuals stages is seen in some Opiliones. For example, abandon the ground; Williams (1962) found Mitopus morio (Fabricius 1799) exhibits ver- individuals of the same species and instar both tical strati®cation, with late instars found at in pitfall traps and on vegetation. Based on high vegetation strata (Adams 1984). Vertical those results, Williams hypothesized opilion- distribution of P. opilio is believed to vary ids may expand their microhabitat distribution 745 746 THE JOURNAL OF ARACHNOLOGY without completely abandoning the ground. rates of conventional pre-emergence herbi- However, it is possible that Williams' (1962) cides (alachlor, metribuzin and chlorimuron results re¯ected diel movement, not microhab- ethyl) and was subsequently treated once itat separation. (2002) or twice (2001) with the post-emer- Opiliones generally are nocturnal (Sankey gence herbicide glyphosate for additional 1949; Todd 1949; Phillipson 1960; Williams weed control. Post-emergence herbicide treat- 1962; Edgar & Yuan 1969). The increase in ments occurred several weeks before trials be- activity at night may be attributed to de- gan. No insecticides were applied during ei- creased light intensity, increased relative hu- ther year. Trials were conducted weekly from midity and decreased temperatures (Todd 23 July±9 September 2001 and from 7±24 Au- 1949). Pfannenstiel & Yeargan (2002), who gust 2002. observed predation events on Helicoverpa zea Phalangium opilio were collected in the (Boddie 1850) eggs in soybean ®elds at 3 h ®eld no more than 5 d prior to the trial dates. intervals during 24 h cycles, found that all ob- Individuals were taken to the laboratory, mea- served events of predation by Phalangiidae sured and/or sexed in order to be placed in a occurred at night. Although P. opilio occa- category (described below), and provided with sionally is active under diurnal conditions, in- food (i.e., H. zea eggs and cornmeal/bacon dividuals exhibit 90% of their total activity diet) and water; food was removed 24 h prior between 1800±0600 h (Edgar & Yuan 1969; to the initiation of observations and individ- Edgar 1980). uals were marked with a small dot of paint 12 Phalangium opilio is known to feed, pri- h prior to the initiation of ®eld observations. marily nocturnally, upon a variety of arthro- In the laboratory, individuals were kept in 8.5 pod pests. In Kentucky, this predator appears 3 8.5 cm (diam 3 ht) containers in incubators to overwinter in the egg stage and undergo at 24 6 1 8C (15:9 L:D) with high humidity three generations per year (Newton & Yeargan via open water containers on the ¯oor of the 2002), with the second generation being the incubators. Prior to the initiation of trials, ®eld most relevant to predation in soybean (due to arenas were checked for naturally occurring seasonal timing of this annual crop), where it opilionids that were removed when found. No feeds on H. zea eggs (Pfannenstiel & Yeargan opilionid species other than P. opilio were en- 2002). Other aspects of its ecology relevant to countered in this study. No other potential its role in soybean ®elds are poorly known, prey were removed from or added to an arena. including its diet breadth, its spatial distribu- Three arenas were monitored on each date. In tion in large ®elds, its within-plant/epigeal each arena, three ®eld collected P. opilio of distribution and its diel activity patterns. We the same category marked with pink ¯uores- investigated the diel activity patterns and mi- cent, water-based paint (Apple Barrel Colors, crospatial distribution of P. opilio for nymphal Plaid Enterprises, Inc.) were introduced si- instars three through seven and both adult sex- multaneously on the ground in the center of es. an arena 1 h before observations began. There is no accurate morphological indicator for sex METHODS or instar in P. opilio nymphs; therefore, size This study was done during the summers of categories (hereafter referred to as P. opilio 2001 and 2002 at the University of Ken- categories) based on cephalothorax width tucky's North Farm near Lexington, KY. In were used (after Newton & Yeargan 2002). each year, three small plots (1 m of soybean Small nymphs were less than 1.0 mm, medi- row per plot) were established within a 0.6 ha um nymphs ranged from 1.0±1.5 mm, and ®eld of soybeans. The soybean variety used in large nymphs were greater than 1.5 mm in both years was Asgrow 4702 and planting oc- cephalothorax width. Adults were discrimi- curred on 1 May 2001 and 20 May 2002. Each nated from nymphs based on the presence or 1 m plot was surrounded by a fence of gal- absence of a genital opening beneath the oper- vanized sheet metal (20 cm tall, 0.5 m from culum (Sankey & Savory 1974) and adult plants on either side of fence); preliminary males and females were identi®ed based on studies showed P. opilio could not scale this the presence or absence of the sexually di- fence. In both years, the entire soybean ®eld morphic horns on the distal segment of male was treated at planting with recommended chelicerae (Sankey & Savory 1974). ALLARD & YEARGANÐACTIVITY PATTERNS OF PHALANGIUM OPILIO 747 During the trials, 5 min observations were of the Department of Entomology at the Uni- made at each arena at 20 min intervals for 1 versity of Kentucky. h at approximately: 1200, 1500, 1800, 2100, Statistical analysis.ÐIn order to analyze 0000, 0300 and 0600 h (EDT). A red light the microspatial distribution and behaviors for ®lter, which minimized disturbance to the an- the different nymphal and adult P. opilio cat- imals, was used to make nocturnal observa- egories, proportions were calculated for each tions and an ultraviolet light was used to lo- set of three individuals during each 1 h ob- cate individuals only if they could not be servation period. There was a maximum of found with the red light. The ultraviolet light nine observations per arena per hour (i.e., was used for ø10% of the observations. Dur- three individuals times three visits 5 the de- ing each observation, the location of individ- nominator for calculating proportions). These uals (i.e., on ground or plant; if on plant, bot- proportions re¯ected the frequency of obser- tom, middle, or top third of the plant, and vations of P. opilio at a particular place exterior or interior portion of the plant) and (ground, bottom of plant, middle of plant, top the behavior of individuals (i.e., walking, of plant) or engaged in a particular behavior grooming, feeding, stationary, palpating, (stationary, walking, feeding, palpating, drink- drinking) were recorded.
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