Developmental Biology of Dictyna Spp. (Araneae: Dictynidae) in the Laboratory and Field Author(S): G
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Developmental Biology of Dictyna Spp. (Araneae: Dictynidae) in the Laboratory and Field Author(s): G. S. Wheeler, J. P. McCaffrey and J. B. Johnson Source: The American Midland Naturalist , Jan., 1990, Vol. 123, No. 1 (Jan., 1990), pp. 124-134 Published by: The University of Notre Dame Stable URL: https://www.jstor.org/stable/2425765 JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at https://about.jstor.org/terms is collaborating with JSTOR to digitize, preserve and extend access to The American Midland Naturalist This content downloaded from 86.59.13.237 on Thu, 01 Jul 2021 09:54:54 UTC All use subject to https://about.jstor.org/terms Am. Midl. Nat. 123:124-134 Developmental Biology of Dictyna Spp. (Araneae: Dictynidae) in the Laboratory and Field G. S. WHEELER,' J. P. McCAFFREY AND J. B. JOHNSON Department of Plant, Soil and Entomological Sciences, University of Idaho, Moscow 83843 ABSTRACT: The developmental biology of two cribellate spider species, Dictyna coloradensis and D. major, was studied in the laboratory (D. coloradensis) and in the field (both species). Laboratory-reared D. coloradensis matured after 6-10 molts and spent an average of 8-13 days per stadium prior to the sixth stadium. Males required an average of 160 days to reach maturity whereas females required 220 days. Mean carapace widths differed among instars, although their ranges overlapped considerably. Field observations revealed univoltine life cycles for both species, with subadults over- wintering and maturing in late spring. Although size of overwintering individuals was similar at the northern and the one southern site (significantly smaller-sized individuals at the other southern site were attributed to mortality of larger individuals due to pesticide drift), Dictyna coloradensis females were significantly larger at the southern, longer-season sites [3000 growing degree days (GDD)] than at the northern, shorter-season site (1500 GDD). Females were largest when reared in the laboratory compared with all field sites. Spiderling field data suggest that development may be arrested at the southern site, syn- chronizing the population and producing similar-sized overwintering individuals. When reared in the laboratory under the same conditions, however, individuals from both the northern and southern sites developed at the same rate and attained the same adult size. INTRODUCTION The cribellate spider, Dictyna major Gertsch (Araneae: Dictynidae) was reported to be the primary mortality factor of the biological control agent Urophora affinis (Frauenfeld) (Diptera: Tephritidae) on spotted knapweed Centaurea maculosa Lamarck (Asteraceae: Cynareae) in Montana (Story and Anderson, 1978; Story and Nowierski, 1984). Since these initial reports, another dictynid spider, D. coloradensis Chamberlin, has also been found in spotted knapweed-infested habitats of northern Idaho (Wheeler, 1985). Dictyna major and D. coloradensis occur sympatrically in northern Idaho and may pose a threat to the biological control of spotted knapweed in that region. Both dictynid species occur on a number of other introduced noxious weeds, some of which also have been targeted for biological control (e.g., yellow starthistle Centaurea solstitialis L.); therefore, the success of other arthropod biological control agents may also be threatened by these spiders. With the exception of taxonomic and geographic information (Chamberlin and Gertsch, 1958) and behavioral data (Jackson, 1977a, 1977b, 1978, 1979, 1980; Jackson and Smith, 1978; Starr, 1988), little information is available on the biology and ecology of the dictynids of the western U.S. rangelands. Thus, our objective was to compare life history parameters (e.g., number, size and duration of the different life stages) of Dictyna coloradensis under laboratory and field conditions. Field data for D. major were also collected and compared with D. coloradensis field data. I Present address: Department of Entomology & Nematology, IFAS 0711, University of Florida, Gainesville 32611-0711 124 This content downloaded from 86.59.13.237 on Thu, 01 Jul 2021 09:54:54 UTC All use subject to https://about.jstor.org/terms 1990 WHEELER ET AL.: SPIDER DEVELOPMENTAL BIOLOGY 125 METHODS AND MATERIALS Laboratory studies. -Dictyna coloradensis egg sacs and maternal webs containing recently emerged spiderlings were collected during the spring and early summer of 1982 and 1983 from two southern sites: the Pond site, 8.3 km NW of Culdesac, and the Central Grade site, 6.4 km NE of Hatwai, Nez Perce County, Idaho. The sites were located ca. 19 km apart, on S-facing slopes, at ca. 658 and 750 m elevation, respectively, and were dominated by yellow starthistle. Additionally, one northern site was studied at Farragut State Park, 1.6 km E of Athol, Kootenai Co., Idaho. This site was located ca. 240 km N of the southern sites, on level ground, ca. 720 m elevation and was dominated by spotted knapweed. Postembryonic development of Dictyna coloradensis in the laboratory was observed twice daily in petri dishes (60 x 15 mm) lined with moistened black filter paper to facilitate observation of egg development. Spiderlings and adults were reared individually and main- tained in 0.185-liter, plastic, snap-top vials and 0.95 liter jars, respectively. All rearing occurred at 27 + 2 C, 45-55% RH and a 14:10 (L:D) photoperiod. The first instar spiderlings emerge from the egg sac as the first free-living and feeding stadium. Preliminary field studies revealed that primarily the first, and to a lesser extent second, instar Dictyna coloradensis spiderlings (recognized by a distinctly darker dorsal coloration of abdominal setae compared with the first instar), remained aggregated in the maternal web. Thus, the first instar spiderlings were allowed to remain in the maternal web and the second instar spiderlings were transferred to individual vials. The diet of the spiderlings consisted of four prey individuals from a mixture of small, acalypterate Diptera and Hymenoptera adults collected from alfalfa and supplied every other day. As winter approached and field-collected prey became scarce, the natural diet was replaced with two wild and two vestigial winged Drosophila melanogaster (Meigen) (Diptera: Drosophilidae) adults every other day. Spiders were observed daily to record all molts, and size was determined by measuring the carapace width of recently ecdysed individuals (?0.02 mm at 50x). Fifty first and second instar spiderlings were sacrificed and measured in 70% ethanol. Later instar spi- derlings and adults were measured live while restrained within a clear plastic sandwich bag. A subset of the entire reared population of Dictyna coloradensis (144 individuals) was measured during the first four instars; all fifth instar through adult individuals were mea- sured. Field studies.-Dictyna spp. development was studied during 1982-1983 at the three locations previously described. During the field season (April-October) all sites were sampled according to a systematic stratified sampling method ca. weekly by hand collecting 20 Dictyna spp. webs along with the resident spider. Five spiders and their webs were collected (one each at 10-m intervals) along each of four randomly selected 50-m transects. Transect directions were selected by randomly determined compass bearings. The spiders were char- acterized by sex, species (when possible), and carapace width using the methods described above. Sex determination was possible only for adults and penultimate males. Only the adult Dictyna individuals or spiderlings in a web inhabited by an adult could be positively identified to species (W. J. Gertsch, pers. comm.); thus, data regarding third through penultimate instars included both species. Additionally, duff and litter samples were collected during the late autumn-winter of 1982-1983 on 31 October and 21 December 1982 and 26 February 1983 (under 0.15-0.5 m of snow for the later two dates) and processed with Berlese funnels. All statistical analyses were conducted using the Statistical Analysis System (SAS Institute Inc., 1987). Experimental designs and analyses [ANOVA, ANCOVA, Student's t test, x2, This content downloaded from 86.59.13.237 on Thu, 01 Jul 2021 09:54:54 UTC All use subject to https://about.jstor.org/terms 126 THE AMERICAN MIDLAND NATURALIST 123(1) Tukey's HSD (P = 0.05), simple linear regression and homogeneity of slopes] followed Sokal and Rohlf (1981). Degrees of freedom for the Student's t test were determined by the Satterthwaite's procedure for unequal sample sizes. When least square mean (Searle et al., 1980) multiple comparisons were conducted, the Bonferroni inequality was used to maintain the 5% experiment-wise significance level. All data are presented as means with the standard error of the mean in parentheses. Growing degree days (GDD) were calculated for the field data by the modified sign-wave method (Allen, 1976) and for the laboratory data by summing the daily degrees between the assumed developmental thresholds of 10 and 35 C. RESULTS Laboratory studies. -All growth parameters were analyzed by site (origin of collection). Only duration of the first instar differed, with spiderlings developing from egg masses collected from the northern site requiring significantly more days (13.1 days, +0.6; n = 117) than those from the southern sites (8.4 days, ?0.8; n = 30) t = 4.8; df = 65; P < 0.0001). Therefore, site data were combined for the remaining analyses. Field-collected adult Dictyna coloradensis spiders maintained in the laboratory deposited an average of 3.0 (?0.2; n = 28) egg sacs per female with each egg mass containing an average of 14.6 (+0.7; n = 66) eggs. Egg clutches were separated by an average of 3.5 days (?0.3; n = 17).