In Alfalfa (Medicago Sativa)
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JEN 127 (2003) J. Appl. Ent. 127, 221–227 (2003) Ó 2003 Blackwell Verlag, Berlin ISSN 0931-2048 Spatial and temporal pattern of colonization of Nabidae (Heteroptera) in alfalfa (Medicago sativa) S. Roth Institute of Ecology, Friedrich-Schiller-University Jena, Germany Ms. received: October 9, 2001; accepted: August 26, 2002 Abstract: Six species of Nabidae (Heteroptera) were collected by standardized sweep net sampling in alfalfa fields in Thuringia, Germany, from 1993 to 1995: Nabis pseudoferus, N. ferus, N. brevis, N. major, Nabicula flavomarginata and Aptus mirmicoides. Colonization of a newly cultivated field was studied over a 3-year period. The density of all the studied nabid species was low (less than five individuals per 100 sweeps) and not related to time since colonization started, or to the distance from the margin of the field. Macropterous species were able to colonize the whole field within one season. The density of one macropterous species, N. pseudoferus, varied between the years of study and was mainly affected by the harvest regime. The brachypterous species reached the margin within one season but for density it took three seasons to reach satiated values also in the centre of the field. The abundance of the brachypterous N. brevis was significantly different both between years and sampling sites. This indicates the importance of the surroundings on the succession of this species. Nabis major, a fully winged species, showed a migration pattern intermediate to macropterous and brachypterous nabids. These results suggest that the total abundance of nabid predators cannot be predicted by time or distance from the expansion source (shelter belts). The abundance of brachypterous nabid individuals can be predicted from time since colonization but is best analysed at the species level. Key words: abundance, flight ability, migration, natural enemies 1 Introduction sition, overall density of nabid predators and density of individual species. Wing reduction occurs in 30% Nabidae are important predators in alfalfa, partic- of the European species of Nabidae (Lattin, 1989). Pimentel ularly on several Aphidae species ( and Because this may be important for the colonization Wheeler Neuenschwander Guppy , 1973; et al., 1975; , pattern, I examined the effect of flight ability separately Siddique Chapman Braman Yeargan 1986; and , 1987; and , for brachypterous and macropterous species. Any Elliot Kieckhefer 1990; and , 1990). The majority of these prediction of the abundance above the species level studies were restricted to American species but there are would considerably facilitate the monitoring of these Boness some studies concerning European species ( , natural enemies of pest species. A priori, I expected both Balogh Loska Geiler Bjegovic 1953; and , 1956; , 1963; , the number of species and their abundances of brac- Balarin Krotova 1968; , 1980; , 1992). A general review of hypterous species to show a decrease with increasing investigations of Nabidae in alfalfa and other crops is distance from refuge zones or field margins and an Lattin given by (1989). increase with increasing time available for coloniza- For biological pest control, it would be desirable to tion. No such dependence was expected for macropter- predict the abundance pattern of these important ous nabids since they can arrive at any point within the predators. For example, the immigration of predators crop. into crops from surrounding shelterbelts or semi- natural habitats affects pest control (Paoletti et al., 1997; Bowie et al., 1999) for several entomophagous arthropods including Carabidae, Staphylinidae, Cocc- 2 Material and methods inelidae and Araneae (Ro¨ser, 1988). For the Nabidae, there was an increase in the abundance in herbicide- 2.1 Sites and time of investigation Mommertz free strips of wheat fields ( , 1993). In The sample site was located in central Thuringia, Germany, Katz addition, et al. (1989) showed that isolated and included an alfalfa field (150 m · 100 m) that was agricultural areas were colonized by aphidophagous bordered by a meadow (150 m · 40 m), field paths and bare, arthropods including Nabidae within one summer. ploughed, land without any vegetation (fig. 1). The alfalfa In this study, the immigration of Nabidae into an field was subdivided into marginal (site M) and central (site alfalfa field was analysed for 3 years after the field’s C) sections with the adjacent meadow designated as a further initial seeding by recording changes in species compo- site (site AM). U. S. Copyright Clearance Center Code Statement: 0931–2048/2003/2704–0221 $ 15.00/0 www.blackwell.de/synergy 222 S. Roth Unused, ploughed land 2.2 Sampling Adult Nabidae were sampled using a standard sweep net (diameter 30 cm) at 3 · 100 sweeps per date and site. In Alfalfa Field grassland habitats, this semi-quantitative method allows X Center (C) comparisons between sites (Roth, 1999). 100 m Standard sweeping was carried out on eight sampling days at intervals of about 10 days between the end of July X Margin (M) and October in 1993 and 1994. In 1995, sampling was carried out on five occasions in August and September because of harvesting in June and September. Sampling Meadow X AM 40 m was not resumed after the second harvest in September 1995. Road 2.3Species 150 m Aspects of general taxonomy and biology of the species are given in Southwood and Leston (1959), Pericart (1987) and Fig. 1. Sample sites (·) in the investigated area with Lattin (1989). Nabicula flavomarginata and Nabis major hibernate in the egg stage, the remaining species as adults the surrounding environment. The distance between Pericart sample site AM (adjacent meadow) and site M (margin ( , 1987). of the alfalfa field) was about 30 m, and the distance between sites M and C (centre of the alfalfa field) was 2.4 Data analysis about 50 m The numbers of nabid species and individuals were compared with regard to sampling site (see fig. 1), the distances between Table 1. Cover of the most dominant plant species in sample sites and different years of the investigation. the margin (site M) and the centre (site C) of the The data were stepwise analysed with regard to: alfalfa field during 1993–1995 using the Braun–Blanquet • Nabidae as group (by pooling all species); scale • wing morphology of the species; • individual species Site M Site C anovass based on Generalized Linear Models (by using SPSS Species 1993 1994 1995 1993 1994 1995 10.0), were used to detect whether factor ÔtimeÕ (year of study) and factor ÔdistanceÕ (by comparing sampling sites) have an Medicago sativa 532555 effect on the number of sampled nabids. In order to fit to Arctium tomentosa r4 –+r normal distribution, data were log-transformed. Agropyron repens +21––– Artemisia vulgaris –+– – r – Convolvulus arvensis r1+–1– Dactylis glomerta –13––– 3Results 3.1 Abundance of all nabids pooled Investigations were conducted from July 1993 until Octo- ber 1995. The field was cultivated with alfalfa (Medicago Six species of Nabidae were caught. Their abundances sativa) in early spring 1993 and first cut in late November are listed in table 2. 1994, with subsequent cuts in June and September 1995. The As shown in table 2, there was no effect of time (by adjacent meadow (AM) was harvested at the same time as the comparing samplings of the years) and distance (by field but it had also been harvested in both late spring and comparing sampling sites) or its interaction on the autumn 1992. The management of the harvest regime of all total number of Nabidae by using anova (F ¼ 1.007, sample sites was determined by the farmer using the land, but no fertilizer and pesticides were used during the time of the d.f. ¼ 8, P ¼ 0.394). study. Only a few individuals of plant species other than alfalfa 3.2 Abundance pooled according to wing morphology were found in sites M and C in 1993. In 1994, the vegetation of site M changed remarkably due to lack of harvesting as The proportion of macropterous and brachypterous revealed by the species inventory for sites C and M (table 1). individuals was not different between the adjacent The vegetation structure of the alfalfa crop also changed sample sites M and C in 1993 and 1995 (the year markedly as a result of the absence of harvesting in the spring of frequent harvesting) (1993: v2 ¼ 1.45, d.f. ¼ 1, n.s.; and summer of 1994. Whereas during 1993 and 1995 the 1995: v2 ¼ 1.21, d.f. ¼ 1, n.s.). In 1994, the total vegetation was about 50-cm high, predominantly vertical in number of macropterous individuals decreased in the structure and with a large number of fresh leaves, during alfalfa field. In addition, the number of brachypterous 1994 the lowest 30–40 cm of vegetation became a dense layer of dry alfalfa stems with a dominantly horizontal structure. individuals increased at the same time at the margin of Fresh leaves and stems were restricted to the upper the field. As a result, the proportion of brachypterous 20–30 cm. The meadow (AM) was dominated by Arrhen- to macropterous individuals changed significantly 2 atherum elatius, Geranium pratense, Poa pratensis, Achillea between sites M and C (1994: v ¼ 7.10, d.f. ¼ 1, millefolium and Falcaria vulgaris. P < 0.01). Colonization of Nabidae in alfalfa 223 Table 2. Number of nabid species sampled and number of all macropterous and brachypterous species pooled in adjacent meadow (AM), margin (M) and centre (C) of alfalfa field in the 3 years under investigation 1993 1994 1995 M + C, in % M + C, in % Total Species AM M C of samples AM M C of samples AM M C number Aptus mirmicoides (BS) 1 1 0 28 0 3 2 38 0 0 0 7 Nabis brevis (BS) 8 8 3 71 4 17 2 100 14 15 13 88 Nabicula Flavomarginata (MS)000 0 000 25012 3 Nabis major (MS) 1 1 0 28 2 1 5 63 0 4 3 17 Nabis ferus (MS) 1 4 4 43 1 2 6 38 2 0 2 22 Nabis pseudoferus (MS) 16 11 12 85 4 7 3 68 7 4 6 70 All macropterous 18 16 16 7 10 14 9 9 13 112 All brachypterous 9 9 3 4 20 4 14 15 13 95 Total number 27 25 19 22 30 28 46 24 26 414 The occurrence of nabid species in alfalfa field (data of sites M and C combined) from 1993 and 1994 is expressed as percentage presence of the eight samples per year.