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Coexistence of Desert Scorpions by Differential Habitat Preference

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Coexistence of Desert Scorpions by Differential Habitat Preference • Coexistence of Desert Scorpions by Differential Habitat Preference STANLEY C. WILLIAMS Reprinted from THE PAN-PACIF1C ENTOMOLOGIST Vol. 46, October, 1970, No. 4 pp. 254-267 Mode in United States of America Coexistence of Desert Scorpions by Differential Habitat Preference STANLEY C. WILLIAMS Department of Biology, San Francisco State College, California 94132 The scorpion fauna of Arizona is rich both in abundance and in number of species. About 19 species have been reported from Arizona in the literature, and these represent four of the six living families in the order Scorpionida. Most of the published research on scorpions has been concerned with systematics, behavior and venoms. Ecological research has been con- spicuously lacking in the literature, with population and community studies being especially uncommon. Baerg (1954) recognized popula- tion periodicity in relation to reproduction and seasonal activity while observing Jamaican scorpions. Alexander and Ewer (1958) discussed the use of temperature-adaptive behavior. Sreenivasa-Reddy (1959) discussed the function of the pectines, and analyzed pectine morphology in relation to burrowing and humidity. Cloudsley-Thompson (1956, 1961, 1962, 1964) discussed biological rhythms, heat death and adapta- tion to desert environments. Gertsch and Allred (1965) made an eco- logical contribution by showing habitat affinities, relative abundances and sex ratios of nine species occurring on the Nevada Test Site. McAlister (1966) discussed the aggregating tendency of Centruroides vittatus (Say) and Smith (1966) discussed the life history of Urodacus abruptus Pocock with emphasis on its home sites. Williams (1966) dis- cussed the obligate burrowing activities of Anuroctonus phaeodactylus, and related this to its ecology. Williams and Hadley (1967) and Hadley and Williams (1968) discussed the habitat preferences, community compositions and nocturnal activities of some populations occurring in Sonora, Mexico. The purposes of this project were two-fold. The first was to study the influence of several different habitat types on the abundance, dis- tribution and community structure of scorpions. The second was to analyze the changes in scorpion distribution and abundance which occur from year to year. The field work was carried out at Phoenix South Mountain, Maricopa County, Arizona from September 1964, to January 1967. Special thanks are due to Mont A. Cazier, David I. Rasmussen, Wendell L. Minckley, and Frank F. Hasbrouck of the Department of Zoology, Arizona State University, for their advice and suggestions. THE PAN-PACIFIC ENTOMOLOGIST 46: 254-267. October 1970 OCTOBER 19701 WILLIAMS-COEXISTENCE OF DESERT SCORPIONS 255 Thanks also are due William S. Parker for assistance in the field, to Charlene F. Williams, Christie Steketee, and William Azevedo for cleri- cal help. This project was partially supported by the National Science Foundation through research grant GB 7679, and by the National Institutes of Health through predoctoral fellowship number 5-F1-GM-23, 794-02. MATERIALS AND METHODS An area approximately three square kilometers in size, located at the eastern end of Phoenix South Mountain (30° 26' N. Lat., 112° 01' W. Long.), Maricopa County, Arizona, was selected for the field study. This area begins about one kilometer west of the town of Guadalupe, along Guadalupe Road. Selection of this study area involved the follow- ing considerations: the ecology was relatively undisturbed; access was convenient; several distinct desert habitats of interest were represented in close proximity; and, the scorpion fauna was abundant and diverse. The general study area was composed of four subareas, each repre- senting a distinct ecological situation. These subareas differed from one another in topography, soil composition, slope, plant community and sun exposure. Essentially, they represented the following ecological situations: an open desert basin near the base of a mountain; a moun- tain canyon slope with a north-facing exposure; a mountain canyon floor; and a mountain canyon slope with a south-facing exposure. Sampling was carried out exclusively by means of unbaited pitfall traps, which were in place for the duration of the study. Number 10 food cans served as traps. These were deep enough to prevent the escape of even the largest scorpion species, and to retain the captured scorpions well below the environmental extremes occurring on the soil surface. Each can was buried so that its open end was flush with the ground surface, and was covered by a suitably sized rock. The trapping method employed has been described in detail by Williams (1968a). A sampling scheme was designed so the four subareas studied could be compared by equivalent trap catch units. For this purpose, the pooled catch of twenty traps was used as a sample unit. Twenty traps were, therefore, established in each subarea. In addition, the data from all subareas could be pooled together to give one sample unit of 80 traps which would reflect the total scorpion activity of the whole area during any given time interval. Because of the seasonal periodicity of the populations in surface activity, the data for trapping samples were pooled for analysis, by 52-weeks. All data were gathered during 104 consecutive weeks from 29 December 1964 through 25 December 1966. 256 THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4 The assumption was made that relative catch numbers were an index of relative abundance for Hadrurus arizonensis (Ewing), Vejovis con- fusus Stahnke and Vejovis spinigerus (Wood). Traps were checked at seven day intervals throughout the year to preserve the equivalence of catch units. The pattern of trap distribution employed was a line-transect along which the traps were placed at vary- ing intervals but never closer than 10 meters apart. Scorpions were removed and preserved (Williams, 1968b) for future reference. Throughout the study, trap maintenance was carried out as needed. This maintenance included removal of soil and animals from traps, re- moval of water after rain, keeping the lip of the trap flush with the soil surface, replacement of traps as they became rusty, and keeping trap sides clean and smooth in order to prevent escape. In addition to the 80 traps sampling the subareas, 100 additional traps were established in other similar areas to test the effect of scorpion removal upon subsequent catches and to test the influence of trap covers on catches (Williams, 1968a). RESULTS DESCRIPTION OF STUDY AREA. Phoenix Mountain is a long, northeast to southwest oriented mountain range, isolated from neighboring moun- tains by expanses of flat desert basin. At the eastern foot of the moun- tain the elevation is 382 meters above sea level while the highest peak reaches an elevation of 758 meters above sea level. The mountain is rugged, rocky and dissected by many irregular valleys and canyons. The mountain belongs to the Basin and Range Physiographic Province of Arizona and is of heterogeneous geologic structure and origin. Most of the eastern end of the mountain is composed of metasedimentary rock of Precambrian origin represented by gneiss and schist associated with dikes of metamorphosed basalt. The remainder of the surface is domi- nated by broad felsic dikes of Tertiary origin and subangular, poorly indurated gravel Quaternary origin (Avedisian, 1966) . The soils of the mountain slopes are generally shallow while those of the open desert surrounding the mountain are deeper, often stony and with a relatively water-impervious, subsurface mineral layer. Climatic studies (Kangieser, 1966) provide the following data for the past 28 years. Air temperature extremes during this period ranged from —13° to 46° C with the lowest temperatures in December and January and highest temperatures during June, July and August. Rain- fall was very irregular from year to year, but generally was greatest in August, and varied during this month from 0.17 to 14.12 cm with an OCTOBER 1970] WILLIAMS-COEXISTENCE OF DESERT SCORPIONS 257 average of 2.84 cm. May and June were the months of lowest rainfall, with monthly rainfall varying from 0.00 to 2.41 cm. The daily high temperature coincided with the occurrence of the daily low relative humidity, this point occurring at about 4:00 p.m. (MST) during the summers. The daily low temperature coincided with the occurrence of the daily high relative humidity, this point being reached at about 6:00 a.m. (MST) during the summers. A number of distinct environmental situations occur because of the physical influence of the mountain. As the mountain is ascended, a heterogeneous array of habitats differentiated by various sun exposures, degrees of slope, water drainages, soil types and plant associations occur. Ecotone effects occur as judged by plant heterogeneity, where two habi- tat types intergrade. Such ecotones are apparent in the canyon bottoms and where the mountain grades into open desert. To study how the various habitats existing at South Mountain affect the distribution and abundance of scorpion populations, four distinct habitats were sampled for comparison. These are described below. Open desert basin.—This area was characterized by a Larrea-Franseria community on open, relatively flat desert (Fig. 1). Since the plant com- munity was more diverse and complex than would have been expected, this area perhaps showed some ecotone effect. The site of this study area was one kilometer west of the town of Guadalupe, along Guadalupe Road. The terrain was basically flat but was occasionally divided by shallow washes. The soils had a texture similar to sandy loam and were well compacted. In most places a hard, subsurface, mineralized layer was present. The vegetation was dominated by creosote bush (Larrea tridentata (D.C.) Coville) and bur-sage (Franseria deltoidea Torr.) , but, paloverde (Cercidium microphyllum (Torr.) Rose and Johnston), ironwood (Olneya tesota Gray), white bur-sage (Franseria dumosa Gray), Mor- mon tea (Ephedra sp.) and saguaro (Carnegiea gigantea (Engelm.) Britt. and Rose) also occurred, although in less abundance. North-facing slope.—This study area was located 1.5 kilometers west of the open desert basin study area, and was a canyon side with a due north exposure (Fig.
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