G)L ! The Journal of ARACHNOLOGY OFFICIAL ORGAN OF THE AMERICAN ARACHNOLOGICAL SOCIETY VOLUME 32 2004 NUMBER 2 THE JOURNAL OF ARACHNOLOGY EDITOR-IN-CHIEF: Daniel J. 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Publication date: 8 November 2004 @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 2004. The Journal of Arachnology 32:193-207 LIFE HISTORIES OF FOUR SPECIES OF SCORPION IN THREE FAMILIES (BUTHIDAE, DIPLOCENTRIDAE, VAEJOVIDAE) FROM ARIZONA AND NEW MEXICO Christopher A. Brown: Department of Biology, Box 5063, Tennessee Technological University, Cookeville, Tennessee 38505 USA. E-mail: cabrown@tntech.edu ABSTRACT* Although scorpions are common and potentially ecologically important members of arid ecosystems throughout the world, basic life history information is lacking for most species. In the current study I examined reproductive investment patterns in four species of scorpion {Centruroides exilicauda, Vaejovis ^spinigerus, Diplocentrus peloncillensis, and Pseudouroctonus apacheanus) from southeastern Arizona and southwestern New Mexico during 1996-1998. Vaejovis spinigerus invested more in repro- duction, in both absolute (total litter mass, TLM) and relative (TLM divided by female mass) terms, than did the other species, and produced the largest litters. Offspring of D. peloncillensis were the largest, weighing over twice as much as the next largest juveniles. Female size was uncorrelated with offspring size in any species, but positive correlations were found between female size and both litter size and total litter mass for C exilicauda (marginally significant) and V. spinigerus (after removal of an outlier). Greater reproductive investment, measured as TLM, was used to make more offspring (in all species but P. apacheanus) but not larger offspring. A marginally significant trade-off between offspring size and number was found in V. spinigerus', there was no size-number trade-off in the other three species. Overall, then, my results suggest that ( 1 ) larger females do not produce larger offspring, (2) larger females may produce more offspring and invest more into a reproductive bout, and (3) the allocation strategy of these species appears to be to invest reproductive resources into production of as many offspring as possible of a relatively fixed size. Keywords: reproductive investment, scorpions, offspring size, litter size, trade-offs One of the primary goals of life history studies is to understand how females allocate energy into reproduction, both within a single reproductive event and across their lifespan (Roff 1992, 2002; Stearns 1992). A female's allocation strategy for a single clutch may be thought of in terms of two “decisions." First, she must choose how much of her available resources to devote to reproduction (i.e., re- productive effort or investment). Second, she must decide how to allocate that resource frac- tion into offspring (i.e., per-offspring invest- ment; Bernardo 1996). The latter decision is generally modeled as a trade-off between making a few large or many small offspring, and a number of studies in various taxa have demonstrated such a trade-off (reviews in Roff 1992; Stearns 1992). Reproductive effort and per-offspring investment have usually been assumed to evolve independently (e.g., Smith & Fretwell 1974; Roff 1992, 2002; Stearns 1992), although recent theoretical and empir- ical evidence suggest they are likely linked evolutioearily (Winkler & Wallin 1987; Caley et al. 2001). Thus, reproductive effort, litter size, and offspring size potentially covary at the phenotypic level. Furthermore, these three traits often vary with female size, which itself is often under strong selective pressure (Roff 1992, 2002; Steams 1992). Based on the above, reproductive allocation patterns are best understood when multiple traits are measured for each of a number of females within a species. For scorpions, such studies are unfortunately rare (Francke 1981; Bradley 1984; Benton 1991a, b; Formanowicz & Shaffer 1993; Brown & Formanowicz 1995, 1996; Lourengo et al. 1996). These studies indicate that, with a few exceptions, female size is unrelated to offspring size. However, larger females generally produce larger litters and have a greater reproductive investment, measured as total litter mass, than smaller females, although these trends do not hold for all species or even all populations of a single species (Brown 2001). Females with greater investment most often simply increase the number of offspring produced, although in 193 194 THE JOURNAL OF ARACHNOLOGY Table 1. — Descriptive statistics (mean ± SE) for life history traits in Centruroides exilicauda, Vaejovis spinigeriis, Diplocentrus peloncillensis, and Pseudouroctonus apacheanus from Arizona and New Mexico. CL = carapace length. CV = coefficient of variation. A dash indicates that a variable was unmeasured. Masses are in mg. Carapace lengths are in mm. Instar 1 duration is in days. Female mass Female CL Mean offspring mass Mean offspring CL Litter size 1996 C. exilicauda 425.2 ±16.1 4.75 ± 0.06 9.4 ± 0.3 1.57 ± 0.02 16.4 ± 1.4 1997 C. exilicauda 373.3 ± 9.7 4.79 ± 0.03 10.3 ± 0.4 — 12.7 ± 1.1 1996 V. spinigerus 882.8 ± 96.1 6.41 ± 0.21 9.8 ± 0.5 — 49.2 ± 4.6 All V. spinigerus 828.4 ± 64.8 6.39 ± 0.14 9.4 ± 0.4 — 48.4 ± 4.0 D. peloncillensis 897.2 ± 57.9 5.42 ± 0.09 23.8 ± 1.5 — 13.0 ± 1.1 P. apacheanus 153.4 ± 17.9 3.66 ±0.13 1.6 ± 0.2 — 27.8 ± 4.2 some cases (Formanowicz & Shaffer 1993; Brown & Formanowicz 1995) larger offspring are also made. Finally, most species do not exhibit an offspring size-number trade-off, and for those that do the strength and direction of the trade-off can vary among populations or years (Brown 2001). The life history of scorpions is virtually unique among terrestrial arthropods (Polis & Sissom 1990). They are often long-lived and relatively large at maturity. Females give birth to live young, potentially producing multiple litters over a number of years, and provide parental care through at least the first molt. Scorpions are also potentially ecologically im- portant predators in many arid and tropical ecosystems (Polis 2001), and knowledge of their life histories should help us explain their ecological effects in these habitats. In this pa- per I report reproductive data, collected in 1996-1998, on four species of scorpion from southwestern Arizona and southeastern New Mexico: one buthid, Centruroides exilicauda (Wood 1863); one diplocentrid, Diplocentrus peloncillensis Francke 1975; and two vaejov- ids, Pseudouroctonus apacheanus (Gertsch & Soleglad 1972) and Vaejovis spinigerus (Wood 1863). For all, I examined relation- ships among female size, offspring size, litter size and reproductive investment. I also ex- amined coefficients of variation in offspring size and their relationship to these traits, since offspring size variation may itself be under selection in certain conditions (Kaplan & Cooper 1984; McGinley et ak 1987). METHODS Study sites and natural history. — Female scoipions were collected from three sites dur- ing 1996 (19-28 May), 1997 (29 June-7 July), and 1998 (21-30 May). All D. peloncillensis and C. exilicauda were collected from Geron- imo Pass (elevation 1770 m) in the Peloncillo Mountains, Hidalgo County, New Mexico, ap- proximately 59 km ENE of Douglas, Arizona (hereafter the Geronimo Pass population. Site 1). All P. apacheanus and some V. spinigerus were collected in the vicinity of the South- western Research Station of the American Mu- seum of Natural History, located in the Chiri- cahua Mountains southwest of Portal, Cochise County, Arizona (hereafter the SWRS popula- tion, Site 2). Elevations ranged from 1620- 1800 m. The remainder of the V. spinigerus were collected from a stretch of Portal Road (elevation 1370-1420 m) approximately 2-5 km east of Portal (hereafter the Portal popula- tion, Site 3). Sites 1 and 2 are primarily Mad- rean evergreen woodland [Brown 1994a; see Erancke (1975) for a further description of the Geronimo Pass site], while site 3 is semidesert grassland (Brown 1994b). Records from SWRS show mean annual precipitation from 1978-1996 was 571 mm, with the wettest months being July and August and the driest months April and May; Geronimo Pass likely shows a similar pattern (Brown 1994a). Scorpions were collected from under rocks or other surface debris during day searches, or while active on the surface at night by using portable flashlights equipped with ultraviolet bulbs. All females were gravid when collected except for seven C exilicauda found in 1997 with first {n = 5) or second {n = 2) instars on the back. Of the four species studied, D. pe- loncillensis is the only obligate buiTower, and was found under rocks at or near the bun'ow BROWN— REPRODUCTION IN SOUTHWESTERN U=S„ SCORPIONS 195 Table 1. — Extended. Total litter mass Relative litter mass CV of offspring mass Instar 1 duration % Survival 153.2 ± 13.3 0.37 ± 0.03 10.1 ± 1.9 7.4 ± 0.2 76.4 ± 7.0 130.7 ± 13.3 0.35 ± 0.04 7.9 ± 1.0 7.2 ± 0.6 89.7 ± 4.9 472.9 ± 46.1 0.56 ± 0.04 9.1 ± 1.1 8.6 ± 0.6 99.7 ± 0.3 440.7 ± 35.7 0.55 + 0.04 9.6 ± 1.0 8.9 ± 0.4 98.7 ± 1.0 310.0 ± 35.9 0.35 ± 0.03 10.4 ± 1.4 13.3 ± 0.8 92.8 ± 3.0 44.1 ± 8.3 0.30 ± 0.05 11.2 ± 3.3 7.8 ± 0.5 100 ± 0 entrance, Vaejovis spinigerus is also known to burrow, and at my collecting sites was captured under rocks with and without obvious burrows. The remaining two species inhabit depressions under rocks, Pseudouroctonus apacheanus was found only under rocks containing moist soil, and was never captured at night. Conversely, C exilicauda and V. spinigerus did not exhibit any noticeable moisture preference and were captured in both day and night searches. Voucher specimens of all species have been de^ posited at the Denver Museum of Nature and Science. Maintenance and data collection. — Fol- lowing capture, females were returned to a laboratory at the University of Texas at Ar- lington, where they were housed individually in 18,5 X 7,5 x 9 cm plastic containers filled with —0.5 cm. of sand. I placed a crumpled paper towel in each container to serve as a refuge; this was kept moistened to increase humidity levels and replaced if it became moldy. I offered each female one adult cricket, Acheta domestica (Linnaeus 1758), every third week while gravid (1-2 juvenile crickets for P. apacheanus); females carrying off- spring were not fed. The laboratory was kept on a 14:10 h lightidark cycle at a mean tem- perature of 26 °C (range 24—31 °C). Addition- al heat was provided by heat lamps (100 W incandescent bulbs with a parabolic metal flashing) attached above the shelves holding the plastic containers. I rotated containers dai- ly along the lengths of the shelves to minimize potential effects of temperature variation on embryo development. Previous work (Brown 1998) has demonstrated that using this addi- tional heat source increases parturition success of females and offspring survival to dispersal in the laboratory. Following birth, first instar juveniles climb onto the female’s back, where they undergo their first molt and subsequently disperse. I therefore checked containers daily for the presence of newborns, newly molted second instars or dispersed offspring. For each litter, all offspring molted or dispersed within a sin- gle 24 h period. Immediately following dis- persal, I weighed the female and all live off- spring individually to the nearest 0.1 mg on an analytical balance (Denver Instruments M- 220). Litter size equaled the number of living and dead juveniles. Females were then killed by heat shock and preserved in 75% ethanol, after which I measured carapace length (CL), to the nearest 0.1 mm, using a dissecting mi- croscope (American Optical) equipped with an optical micrometer. Using the same pro- cedure I also measured offspring CL on a sub- set of C. exilicauda litters from 1996. For my measure of reproductive investment I calculated total litter mass (TLM) by sum- ming individual offspring masses. I also cal- culated a size-corrected measure of invest- ment, relative litter mass (RLM), as TLM divided by female mass. Both TLM and RLM will be underestimated by using masses of second, rather than first, instars, as scorpions lose weight between birth and dispersal (For- manowicz & Shaffer 1993). This is more like- ly to affect interspecific comparisons since mass loss rates probably vary more among than within species, although the degree to which this is true is unknown. As a measure of within-litter variation in offspring mass I calculated coefficients of variation (CVs) us- 196 THE JOURNAL OF ARACHNOLOGY ing the bias correction of Sokal & Rohlf (1995:58). Finally I calculated percent surviv- al of offspring until dispersal as the number of juveniles alive at the time of weighing di- vided by litter size. Data analyses. — Because they have more space in which to store embryos or are better at obtaining resources, larger females often are predicted to produce more or larger off- spring and to have greater reproductive in- vestment. Thus, I examined relationships be- tween female size (CL; see Results) and offspring size, litter size, or TLM using least- squares regression. For the remaining relation- ships I calculated correlation coefficients (Pearson’s r), as I had no a priori basis for cause and effect. The trade-off between off- spring size and number was examined in the following manner to statistically control for variation in female size. I first regressed off- spring mass or litter size against female CL and obtained residuals. I then used these re- siduals in a correlation analysis. For each spe- cies separately, I initially set a = 0.05 and then made adjustments using the sequential Bonferroni procedure (Rice 1989) to account for multiple (n = 10) significance tests. Re- gression and correlation results were com- bined within species for this correction. When P values were less than 0.05 but nonsignifi- cant after Bonferroni correction, I also report adjusted a values. For C exilicauda I collected sufficient data to make comparisons between years using analysis of variance. Because of statistical problems associated with ratios (Packard & Boardman 1987), in this ANOVA I used the residuals from a regression of TLM on female mass as a measure of RLM. For all analyses reproductive variables were logiy-transformed to meet assumptions of parametric tests. Data analysis was carried out using Statistica for Windows version 4.5 (StatSoft 1993). RESULTS I obtained data from 63 females, as follows: 33 C. exilicauda (1996: n = 18; 1997: n = 15), 19 K spinigerus (1996 SWRS: n = 3; 1996 Portal: n =. 12; 1997 SWRS: n = 2; 1997 Portal: n = 2), six D. peloncillensis (1996, n = 4; 1997: n — 2), and five P. apa- cheamis (1996: n = \\ 1997: n = 3; 1998: n = 1). Since relationships among reproductive variables can vary over space or time (e.g.. Brown & Formanowicz 1995; Brown 2001), it is preferable to use data from a single pop- ulation and breeding season. I have therefore examined each year separately for C. exilicau- da and have calculated two sets of means for V. spinigerus, one using all data and one using only 1996 Portal data (which had the largest sample size). For D. peloncillensis and P. apacheanus I lacked enough individuals in any one year, and so combined data across years. Summary statistics for each species are presented in Table 1. Centruroides exilicauda females gave birth between 13 June-14 July in 1996 and from 1- 9 July in 1997. Parturition in both D. pelon- cillensis (16 August-3 September) and P. apacheanus (29 July-18 August) occurred lat- er in the season. Portal V. spinigerus had the most protracted birthing period, from 30 June- 10 August in 1996 (both 1997 females gave birth in mid July). Vaejovis spinigerus from SWRS gave birth later than Portal fe- males in both 1996 (10-26 August) and 1997 (2-17 August). Diplocentrus peloncillensis and V. spinige- rus were similar in mass and the largest spe- cies in this study, weighing about twice as much as C. exilicauda and 5-6 times more than P. apacheanus. However, these differ- ences in female size were not necessarily re- flected in other life history traits (Table 1). Offspring of D. peloncillensis averaged twice the mass of V. spinigerus offspring, while C. exilicauda and V. spinigerus offspring were similar in mass despite their two-fold differ- ence in adult size. However, V. spinigerus had 3-4 times as many offspring as either D. pe- loncillensis or C. exilicauda. The smallest species, P. apacheanus, also had litters 1.7-2 times larger than D. peloncillensis or C. exil- icauda, albeit with much smaller offspring. Combining offspring size and litter size, V. spinigerus invested more in reproduction than the other species, both in absolute (TLM) and relative (RLM) terms. The remaining species, while differing substantially in total invest- ment, had similar RLM values. The per-litter percentage of offspring surviving until dis- persal was very high (> 90%), with the ex- ception of C. exilicauda in 1996. Dead off- spring were most often first instars, many of which had died while molting. Centruroides exilicauda did not differ be- tween years in female CL (F, 3, = 0.38, P = BROWN— REPRODUCTION IN SOUTHWESTERN U.S. SCORPIONS 197 Female CL (mm) Eigures 1-4. — Linear regressions of mean offspring mass against female carapace length for four species of scorpion from Arizona and New Mexico. Regression equations are given in Table 2. Dashed lines indicate a nonsignificant regression. (1) 1996 (filled circles, heavy line) and 1997 (open circles, light line) Centruroides exilicauda. (2) 1996 Portal (filled circles, heavy line) and all (filled + open circles, light line) Vaejovis spinigerus. The square indicates an outlier (see text). (3) Diplocentrus peloncillensis. (4) Pseudouroctonus apacheanus. 0.54), offspring mass (F, 3i = 2.71, P — 0.11), litter size (Fj 3, - 3.25, P = 0.08), TLM (F^ 31 - 1.21, P = 0.28), RLM (Fi 3, = 0.42, P = 0.52) or within-litter variation in offspring mass (Fi 3i = 1.23, P = 0.28). However, fe- male mass was significantly greater in 1996 (F, 3, = 5.83, P = 0.02). Repeating the above analyses using female mass as the covariate in an ANCOVA again revealed no significant differences among years (results not shown). Female mass and CL were strongly posi- tively correlated in U. spinigerus (1996 Portal: r = 0.87, P < 0.001, n = 12; all females: r = 0.94, P < 0.001, n = 19), P. apacheanus (r = 0.96, P = 0.04, n = 4), and C. exilicauda in 1996 (r = 0.83, P < 0.001, n = 18). The relationship between these variables was also positive, but not significant, for D. peloncil- lensis (r = 0.60, P = 0.21, n = 6) and C. exilicauda in 1997 (r = 0.19, F = 0.49, n = 15). I therefore used CL as my measure of female size because it is less subject to fluc- tuations (e.g., due to feeding history) than is mass. Results using female mass were quali- tatively similar unless otherwise noted. Mass and CL were also significantly positively cor- related for C. exilicauda offspring in 1996 (r = 0.75, P = 0.01, n = 10), and I therefore report only correlations involving mass (using CL gave similar results) to be consistent with the 1997 data. For C. exilicauda, D. peloncillensis, and F. apacheanus, female size was uncorrelated with offspring size, litter size, or total invest- ment following Bonferroni correction (Table 2; Figs. 1, 3-5, 7-9, 11, 12). For C. exilicau- da, marginally significant positive correlations were found between female size and litter size 198 THE JOURNAL OF ARACHNOLOGY Table 2. — Linear regression statistics for the relationship between female size (carapace length) and offspring mass (OM), litter size (LS), or total litter mass (TLM) for four species of scorpion from Arizona and New Mexico. All variables were log-transformed prior to analysis, df = degrees of freedom for the E-test. Species Variable Slope Intercept E2 E E 1996 C. exilicauda OM -1.17 4.06 0.21 4.28 0.055 df = 1, 16 LS 4.10 -3.67 0.31 7.08 0.017 TLM 2.93 0.39 0.15 2.80 0.11 1997 C. exilicauda OM 0.54 1.47 0.007 0.10 0.76 df = 1 , 13 LS 7.62 -9.46 0.32 5.99 0.029 TLM 8.16 -7.99 0.31 5.83 0.031 1996 Portal V. spinigenis OM 0.36 1.61 0.05 0.56 0.47 df = 1, 10 LS 1.16 1.70 0.13 1.56 0.24 TLM 1.51 3.30 0.25 3.32 0.10 All V. spinigerus OM 0.20 1.85 0.01 0.18 0.68 df = 1, 17 LS 1.73 0.60 0.15 2.88 0.11 TLM 1.93 2.45 0.22 4.79 0.042 D. peloncillensis OM 1.11 1.29 0.08 0.33 0.60 df = 1,4 LS 3.86 -3.98 0.47 3.60 0.13 TLM 4.97 -2.68 0.49 3.84 0.12 E. apacheanus OM 2.24 -2.49 0.26 0.70 0.49 df = 1,2 LS -3.64 7.89 0.79 7.51 0.1 1 TLM - 1 .40 5.39 0.21 0.53 0.54 in both years (1996: P — 0.017, adjusted a = 0.0056; 1997: P = 0.029, adjusted a = 0.0056) and between female size and TLM in 1997 {P = 0.031, adjusted a = 0.0063). The regressions involving litter size and TLM for C. exilicaiida in 1997 were in the same direc- tion when female mass was used in place of CL, but were much weaker and not significant (log litter size = 1.4 + 0. 18*log female mass, - 0.002, F, ,3 = 0.03, P = 0.86; log TLM = 2.9 + 0.32*log female mass, = 0.007, F| ,3 = 0.09, P = 0.77). Surprisingly, both lit- ter size and TLM declined with increasing fe- male size in P. apacheanus. Female size was also uncorrelated with re- productive traits in V. spinigenis after Bon- ferroni correction (Table 2; Figs, 2, 6, 10), al- though there was a marginally significant relationship between female size and TLM when using data from all litters {P = 0.042, adjusted a = 0.0063). However, one data point (the square in Figs. 2, 6, 10) was deter- mined to be an outlier. With this removed there was no change in the relationship be- tween female size and offspring mass (results not shown), but both litter size and TLM were now significantly positively correlated with female size (litter size: 1996 Portal: log litter size =—2.13 + 3.28*log female CL, = 0.84, E| 9 = 44.9, P < 0.0001; all females: log litter size = —3.02 + 3.72*log female CL, R^ = 0.48, E, „ = 14.5, P = 0.002; TLM: 1996 Portal: log TLM = 0.41 + 3.12*log female CL, E2 = 0.69, E, 9 = 19.8, P = 0.002; all females: log TLM = -0.05 + 3.31*log fe- male CL, E2 = 0.43, E, ,6 = 12.0, P = 0.003). Reproductive investment, measured as TLM, was uncorrelated with offspring mass in all species (Table 3; Figs. 13-16), although greater investment tended to be associated with larger offspring in D. peloncillensis and E. apacheanus. Offspring number was signif- icantly positively correlated with TLM in C. exilicauda and V. spinigerus, and the corre- lation between these traits was marginally positive in D. peloncillensis {P = 0.04, ad- justed a = 0.005; Table 3; Figs. 17-20). I found no evidence of a trade-off between off- spring size and number (Figs. 21-24) in C. exilicauda (1996: r = 0.21, P = 0.40, n = 18; 1997: r = -0.12, P = 0.68, n = 15), E. apa- cheanus (r = —0.74, E = 0.26, n = 4), or D. peloncillensis (r = —0.18, E = 0.73 n = 6). There was also no trade-off in the 1 996 Portal V. spinigerus (r = —0.45, E = 0.14, n — 12), although a marginally significant trade-off was found when including all data (r = -0.50, E = 0.03, adjusted a = 0.0056, n = BROWN— REPRODUCTION IN SOUTHWESTERN U.S. SCORPIONS 199 4.2 4.4 4.6 '4.8 Female CL (mm) Female CL (mm) 18 .§ 14 12 10 32 ® O 7 30 8 / 28 |26 bo fc 24 22 20 • 18 16 __J 1 L_ ^ ^ 2 5,1 5.2 5.3 5.4 5.5 5.6 5.7 Female CL (mm) 3.4 3.5 3.6 3.7 3.8 Female CL (mm) 3.9 Figures 5-8. — Linear regressions of litter size against female carapace length for four species of scorpion from Arizona and New Mexico. Regression equations are given in Table 2. Short dashed lines indicate a nonsignificant regression, long dashed lines a marginally significant regression. Symbols as in Figure 1. (5) Centruroides exilicauda. (6) Vaejovis spinigerus. (7) Diplocentrus peloncillensis. (8) Pseudoiiroctonus apacheanus. 19). Within-litter variation in offspring mass (CV) was uncorrelated with other traits in most species (Table 3); the only exceptions were marginally negative correlations be- tween CV and litter size {P = 0.044, adjusted a - 0.0063) or TLM {P = 0.026, adjusted a = 0.0056) for the 1996 Portal V. spinigerus. No obvious trends were found in CV patterns across species. Removal of the outlier for V. spinigerus qualitatively altered none of the above results (results not shown). Captivity is suspected to affect scorpion re- production because of differences in prey availability or environmental parameters be- tween the field and laboratory (Polls & Sis- som 1990). Therefore, for each species I ex- amined correlations between days in captivity (the time between capture and parturition) and female mass, offspring mass, litter size, and total litter mass, the traits most likely to be affected by laboratory conditions. Within a species, these traits tended to either all in- crease or all decrease (Table 4), although there was no consistent overall pattern across spe- cies. Only two correlations were marginally significant: litter size of V. spinigerus (all lit- ters; P = 0.05, adjusted a = 0.0125), and mass of female C. exilicauda in 1997 {P = 0.04, adjusted a = 0.0125), declined with time spent in captivity. The results for C exilicau- da are likely unimportant, as all litters had dispersed within 18 days of capture. Both C. exilicauda females captured in 1997 carrying second instar juveniles gave birth in the laboratory to a second litter. The times between dispersal of the first litter and birth of the second were 109 and 341 days. Both females increased substantially (93-94 200 THE JOURNAL OF ARACHNOLOGY Female CL (mm) Female CL (mm) Female CL (mm) Figures 9-12. — Linear regressions of total litter mass against female carapace length for four species of scorpion from Arizona and New Mexico. Regression equations are given in Table 2. Short dashed lines indicate a nonsignificant regression, long dashed lines a marginally significant regression. Symbols as in Figure 1. (9) Centniroides exilicauda. (10) Vaejovis spinigenis. (11) Diplocentrus peloncillensis. (12) Pseudouroctomis apachecmus. mg) in post-dispersal mass and produced larg- er second litters (first litter/second litter: 8/18 and 13/14). Mean offspring mass in the sec- ond litter increased for one female and de- creased for the other (10.1 mg/8.9 mg and 10.5 mg/ 11.8 mg, respectively, for the litter sizes above). Neither female had access to males after capture, but I do not know whether females had remated in the field while gravid or whether sperm from a single mating was used for both litters. DISCUSSION Based on data summarized in Polis & Sis- som (1990) and Brown (2001), V. spinigerus had larger litters than other species in the fam- ily Vaejovidae [mean = 27.5 {n = 22)] or the genus Vaejovis [mean = 29.0 (n = 10)]. Pre- vious authors have reported litter sizes of 13- 69 for V. spinigerus {n = 4 litters; McAlister 1960; Stahnke 1966; Williams 1969); two Portal females had litters larger than this max- imum (70 and 76 juveniles). First instar du- ration was shorter than the family mean of 12.6 d {n = 8), but slightly longer than the duration reported for V. spinigerus by Mc- Alister (1960; 7-8 d). In contrast, both C. exilicauda and D, pe- loncillensis had smaller litters on average than other confamiliars [Buthidae mean = 22.8 (n = 33), Diplocentridae mean — 24.6 (n = 8)] or congeners [Centruroides mean = 37.5 (n = 7), Diplocentrus mean = 24.8 {n = 6)]. Geronimo Pass C. exilicauda had litters inter- mediate in size to those of inland (mean = 10.1) and coastal (mean = 18.2) C. exilicauda from Baja California [Myers 2001; however, this likely represents a distinct species from the New Mexico population (Gantenbein et al. 2001)], but smaller than the mean of 20 for BROWN— REPRODUCTION IN SOUTHWESTERN U.S. SCORPIONS 201 Table 3. — Correlations (Pearson’s r) between reproductive traits in four species of scorpion from Arizona and New Mexico. ECL = female carapace length. All other abbreviations are defined in Tables 1 and 2. * P < 0.05, *** P < 0.001. Correlation between: TLM-OM TLM-LS CV-FCL CV-OM CV-LS CV-TLM 1996 C. exilicauda in = 18) 0.24 Q 94*** 0.37 -0.36 -0.20 -0.31 1997 C. exilicauda {n = 15) 0.38 0.90*** 0.40 -0.07 0.06 0.03 1996 Portal V. spinigerus (n = 12) 0.17 0.88*** -0.29 -0.05 -0.59* -0.64* All V. spinigerus in = 19) 0.004 0 90*** -0.20 -0.34 -0.13 -0.30 D. peloncillensis {n = 6) 0.61 0.83* -0.20 0.04 0.17 0.15 P. apacheanus (n = 5) 0.48 0.71 -0.32 -0.86 0.13 -0.52 this species (Polis & Sissom 1990). First in- star duration was similar to family means for C. exilicauda [mean = 6.5 d (n = 20)] and D. peloncillensis [mean = 14.4 d (n = 4)], and for C. exilicauda was within the range previously reported for this species (6-15 d: Stahnke 1966; Williams 1969). Pseudouroc- tonus life history data are available for only a single litter of P. reddelli (Gertsch & Soleglad 1972). This species is much larger than P. apacheanus and has larger litters and a slight- ly shorter first instar duration (Brown 1997). With the exception of V. spinigerus, relative litter mass was lower than in any sexually re- producing scorpion (range 0.44-0.55: Benton 1991a; Formanowicz & Shaffer 1993; Brown & Formanowicz 1995, 1996; Lourengo et al. 1996). Such low relative investment might oc- cur if females cannibalize some newborns, de- creasing observed litter sizes. However, no fe- males in this study were observed feeding on juveniles. Low food levels might also lead fe- males to invest fewer resources in reproduc- tion, or to resorb some embryos (Polis & Sis- som 1990). If precipitation levels can be used as a proxy for arthropod prey availability, then reduced investment might be expected in 1996, when only 27.4 mm of precipitation had fallen by the end of May when scorpions were collected. However, RLM was as low in 1997 (e.g., for C exilicauda), when > 7 times as much rain (202.4 mm) had fallen in the same period. Conditions in the laboratory might also have been more stressful than in the field, leading to decreased investment in reproduc- tion (lower TLM), maintenance (lower female mass), or both, as time spent in captivity in- creased. If so, low RLM should primarily re- flect declines in TLM, a trend not supported by my data for any species. Thus, relatively low reproductive investment in C. exilicauda, D. peloncillensis, and P. apacheanus may simply reflect an adaptive response by these species to specific environmental conditions, such as a decrease in the length of the growing season or lower prey availability, in lower montane woodlands. Obviously, more com- parative investment data for scorpions from various habitats are required to assess this hy- pothesis. For C. exilicauda and D. peloncillensis, more detailed comparisons can be made to C vittatus (Say 1821) and D. Undo Stockwell & Baldwin 2001. The latter two species occur sympatrically at Chandler Independence Creek Preserve in west Texas (Brown & For- manowicz 1995, 1996), which differs in ele- vation (—700 m) and habitat (Chihuahuan de- sert scrub) from Geronimo Pass. Reproductive data were obtained for the Texas species in 1992 (Brown & Formanowicz 1995, 1996) and 1996-1997 (Brown unpub. data). Diplo- centrus females are similar in size, while C. vittatus females are —30% larger by mass than C. exilicauda females. For the other reproduc- tive traits, the same pattern emerged for each 202 THE JOURNAL OF ARACHNOLOGY Total Litter Mass (mg) Total Litter Mass (mg) Total Litter Mass (mg) Figures 13-16. — Correlations between total litter mass and mean offspring mass for four species of scorpion from Arizona and New Mexico. Correlation coefficients are given in Table 3. Symbols as in Figure 1. (13) Centriiroides exilicciiida. (14) Vaejovis spinigenis. (15) Diplocentnis peloucUlensis. (16) Pseiidouroctoniis apachecuius. congener pair: Texas scorpions had larger lit- ters, smaller offspring, and larger total and rel- ative investment than New Mexico scoipions (part of this trend was relaxed for D. Undo in 1997, as TLM and RLM were just slightly larger than comparable values for D. pelon- ci liens is). The proximate cause of these interspecific differences is unknown, given that the two study sites differ in a number of environmen- tal characteristics. For instance, temperature is known to affect growth rates, adult size, and reproduction in a variety of ectotherms (e.g., Li & Jackson 1996; Ernsting & Isaaks 2000). In particular, theoretical and empirical studies (e.g., Yampolsky & Scheiner 1996; Ernsting & Isaaks 2000) have demonstrated that egg/ offspring size decreases, and litter size in- creases, with an increase in temperature. De- creases in offspring size may not balance increases in offspring number, so that total in- vestment may be greater at higher tempera- tures (Ernsting & Isaaks 2000). Given that mean and maximum monthly temperatures are higher at Independence Creek than Geronimo Pass across the year [using data from the southern (NCDC 2002a) and western (NCDC 2002b) regional climate center websites, re- spectively], the trends for Centruroides and Diplocentrns are in the predicted direction. Alternatively, larger offspring size may be fa- vored at Geronimo Pass if predation pressure on juveniles is greater or food availability for juveniles is lower than at Independence Creek. Under such conditions large offspring are pre- dicted to survive better and/or develop faster than small offspring (Shine 1978; Ito & Iwasa 1981). Female size had little influence on offspring size, but larger females tended to have larger BROWN— REPRODUCTION IN SOUTHWESTERN U.S. SCORPIONS 203 Table 4. — Correlations (Pearson’s r) between days spent in captivity and selected reproductive traits in four species of scorpion from Arizona and New Mexico. All abbreviations are defined in Tables 1 and 2. *F < 0.05. Correlation between days in captivity and: Female mass OM LS TLM 1996 C exilicauda {n = 18) 0.40 -0.14 -0.30 -0.34 1997 C. exilicauda {n = 15) -0.54* 0.17 0.35 0.39 1996 Portal V. spinigerus {n = 12) 0.15 -0.05 -0.44 -0.48 All V. spinigerus in = 19) -0.01 0.33 -0.46* -0.35 D. peloncillensis (n = 6) 0.75 0.47 0.40 0.58 P. apacheanus {n = 5) 0.54 0.71 0.16 0.66 litters and invest more into reproduction in all species but F. apacheanus. The latter two re^ lationships are somewhat equivocal, as the correlations between female size and litter size or TLM were nonsignificant (for D. pelonciF lensis), marginally significant (for C exilicau- da), or significant only after removal of an outlier (for V. spinigerus). Nevertheless, these trends are likely to reflect real and biologically meaningful relationships, given that a positive correlation between female size and litter size or total investment is common in other taxa (reviewed in Roff 1992; Stearns 1992), in- cluding other arachnids (e.g., solifuges: Punzo 1998; spiders: Kessler 1971; Killebrew & Ford 1985; McLay & Hayward 1987; Punzo & Henderson 1999). In a recent review, Brown (2001) examined allometric relation- ships involving offspring size or number for 1 1 species of scorpion (including three of the four current species), some represented by multiple populations or years. Adding two ad- ditional sources (Formanowicz & Shaffer 1993; Myers 2001), significant positive cor- relations with female size (measured as mass in most cases) were found in only six of 30 cases for offspring size (with two additional significant negative correlations) and 12 of 32 cases for litter size (with one significant neg- ative correlation). Total litter mass appears to correlate more strongly with female size for scorpions in general, as in six of eight previ- ous cases this relationship was at least mar- ginally significant (Bradley 1984; Benton 1991b; Formanowicz & Shaffer 1993; Brown & Formanowicz 1995, 1996). Thus, my re- sults agree in general with those from previ- ous studies of scorpion reproduction, although my support for an allometric effect on litter size was stronger than in Brown (2001). In the current study and in Brown (2001), two issues may obscure detection of signifi- cant allometric effects. First, sample sizes may be too low to reveal effects; for example, in the current study the significant or marginally significant results involving litter size and TLM came from the two species (C exilicau- da and V. spinigerus) with the largest sample sizes. The use of Bonferroni correction, al- though conceptually justified, may exacerbate this problem by making it harder to detect im- portant trends. I note, however, that across species of scorpions sample size is uncorre- lated with the magnitude of the correlation co- efficient between female size and litter size [r = 0.005, using data from Brown (2001)]. Sec- ond, and perhaps more importantly, the choice of female size measure appears to alter the strength of the correlation with litter size (and TLM) in a usually consistent way. In the cur- rent study, in all cases litter size was more strongly correlated with female CL (i?^ range without removing V. spinigerus outlier, 0.13- 0.79) than with female mass range, 0.002- 0.12), although only for 1997 C exilicauda did statistical significance change. The same 204 THE JOURNAL OF ARACHNOLOGY Total Litter Mass (mg) Total Litter Mass (mg) Figures 17-20. — Correlations between total litter mass and litter size for four species of scorpion from Arizona and New Mexico. Correlation coefficients are given in Table 3. Symbols as in Figure 1. (17) Centruroides exilicauda. (18) Vaejovis spinigerus. (19) Diplocentrus peloncillensis. (20) Pseudouroctonus apacheamis. trend holds in Tityus columbianus (Thorell 1876), where litter size is strongly correlated with female body length but weakly correlated with female mass (Lourengo et al. 1996), and C vittatus, where litter size and TLM are more strongly correlated with female CL than female mass in seven of ten population-year combinations (Brown 1998). Thus, female mass may be a relatively poor measure for examining allometric relationships in scorpi- ons, and significant correlations between litter size and female size may be more common than reported in Brown (2001). The lack of significant relationships be- tween offspring size and female size or TLM suggests that offspring size is relatively can- alized compared to litter size. This was reflect- ed in coefficients of variation across females; CVs of litter size (range of species means, 21.2-42.5%) were 1.5-3 times greater than CVs of offspring size (range, 14.2-27.6%). My results are consistent with those from oth- er terrestrial ectotherms (e.g., lizards: Cong- don 1989; spiders: Kessler 1971; McLay & Hayward 1987; Killebrew & Ford 1985; in- sects: Mappes et al. 1996; scorpions: Brown & Formanowicz 1995, 1996; Lourengo et al. 1 996) indicating that increases in reproductive output result primarily from adding offspring rather than increasing offspring size. Canali- zation of offspring size may represent a de- cision by females to allocate a relatively con- stant amount of resources to each offspring. As suggested for spiders (Marshall & Gittle- man 1994), this amount may be near the min- imum necessary to ensure survival of off- spring until dispersal. Variation in offspring size among or within litters might then reflect “noise” created by a female’s inability to pre- cisely allocate resources. Alternatively, the BROWN— REPRODUCTION IN SOUTHWESTERN U.S. SCORPIONS 205 Figures 21-24. — Correlations between residual offspring mass and residual litter size for four species of scorpion from Arizona and New Mexico. Correlation coefficients are given in the text. Symbols as in Figure 1. (21) Centruroides exilicauda. (22) Vaejovis spinigerus. (23) Diplocentrus pelonciUensis. (24) Pseudouroctonus apacheanus. relative uniformity in offspring size may re= fleet anatomical constraints (see e.g., Congdon & Gibbons 1987), perhaps in the structure of the ovariuterus or genital operculum, which limits egg or offspring size but is unrelated itself to female size. I found little support for an offspring size- number trade-off in the four species studied, similar to the trend for scorpions in general (Brown 2001). Also similar to results from other arachnids (Simpson 1993; Brown 1998, 2001) was the year-to-year change in strength and magnitude of the trade-off in C. exilicau- da. The reasons for this are unclear, as much life history theory predicts that offspring size and number will be negatively correlated if resources available for reproduction are lim- ited (Roff 1992, 2002; Stearns 1992). Varia- tion in acquisition of resources, which can oc- cur if better quality females obtain more or better quality prey, has been predicted to po- tentially obscure trade-offs between offspring size and number (van Noordwijk & de Jong 1986). Brown (in press) has recently demon- strated that this hypothesis applies to scorpi- ons, that is, that stronger (more significant) negative correlations between offspring size and number occur when variation among fe- males in reproductive investment (a measure of variation in resource acquisition by fe- males) is relatively low. A second explanation for the lack of a trade-off involves the rela- tively constant size of offspring. If there exists a fixed allocation strategy, that is, females use their reproductive resources to make as many offspring as possible of a given (minimal?) size, then there is no underlying basis for a trade-off. In summary, I found little evidence for larg- er females to produce larger offspring for any 206 THE JOURNAL OF ARACHNOLOGY of the species studied. However, for two of the four species (C. exilicaiida and V. spini- genis) larger females produced more offspring and had a greater total litter mass, although these results were not always significant after Bonferroni correction. For each of the four species studied, females with higher reproduc- tive investment produced more but not larger offspring than females with lower investment. Finally, no trade-off existed between offspring size and number, except when I combined data across years and populations in V. spinigems. 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Williams, S.C. 1969. Birth activities of some North American scorpions. Proceedings of the Califor- nia Academy of Sciences 37:1-24. Winkler, D.W. & K. Wallin. 1987. Offspring size and number: a life history model linking effort per offspring and total effort. American Natural- ist 129:708-720. Yampolsky, L.Y. & S.M. Schemer. 1996. Why larg- er offspring at lower temperatures? A demo- graphic approach. American Naturalist 147:86- 100. Manuscript received. 18 October 2002, revised 27 June 2003. 2004. The Journal of Arachnology 32:208-220 POPULATION DYNAMICS OF AN ISOLATED POPULATION OF THE HARVESTMAN ILHAIA CUSPIDATA (OPILIONES, GONYLEPTIDAE), IN ARAUCARIA FOREST (CURITIBA, PARANA, BRAZIL) Luiz Augusto Macedo Mestre: Universidade Federal de Sao Carlos, DEBE, Dep. Ecologia e Biologia Evolutiva, Av. Washington Luis, Sao Carlos SP, Brazil Ricardo Pinto-da-Rocha': Departamento de Zoologia, Institute de Biociencias da Universidade de Sao Paulo, Caixa Postal 11461, 05422-970 Sao Paulo SP, Brazil. E-mail: ricrocha@usp.br ABSTRACT. The harvestman Ilhaia ciispidata was studied in Curitiba, state of Parana, Brazil. The site studied is a 30,000 forest remnant of Araucaria forest surrounded by houses. Harvestmen were sampled 21 times over 14 months (June 1997-August 1998); at intervals of 21-30 days. The population size was estimated by Fisher-Ford and Jolly methods, and did not vary considerably from autumn (June) to spring (November 1997). However, it increased rapidly (300 %) during late spring and summer (December- March). The lowest population size estimated was 1,429 adults/subadults, during the winter (June 1997) and, the highest was 14,445 during the autumn (April 1998). The recapture rates ranged from 16%-41%. The sex ratio observed in all sampling periods was 1:1. The density varied from 0.05 (winter) to 0.47 adult + subadults/m^ (autumn). The extremely different abundances observed between seasons could have been influenced by temperature. The immatures were observed all year, suggesting a continuous repro- duction, but they were much more abundant during spring and summer. Ecological aspects including aggregation, individual movement and life span were also discussed. Keywords: Araucaria forest, Ilhaia cuspidata, Opiliones, Population ecology Harvestmen are normally solitary, noctur- nal, omnivores, vagile and photophobic (Cod- dington et al. 1990; Savory 1938). They are commonly found in humid forests, under fall- en trunks, in leaf litter, mosses and inside caves (Edgar 1990). Members of this order are very common in Neotropical forests, where their diversity can be very high, in the Atlan- tic Rain Forest of Brazil, more than one har- vestmen can be found per square meter (Pinto- da-Rocha pers. obs.). In temperate zones harvestmen biomass sometimes can exceed that of spiders and they may be important con- trollers of insect populations (Hillyard & San- key 1989). Data on population biology and the natural history of Opiliones in South America are very scarce (see Gnaspini 1996; and Pinto-da- Rocha 1999 for references). Specific studies related to the ecology and population biology of Neotropical harvestmen have been con- ' Corresponding author. ducted with Goniosoma spelaeum (Mello-Lei- tao 1933) (Gonyleptidae, Goniosomatinae) in caves of the Ribeira valley, Sao Paulo, Brazil (Gnaspini 1996), Pachylospeleus strinatii Sil- havy 1974 in the “Gruta das Areias de Cima”, Sao Paulo, Brazil (Pinto-da-Rocha 1996a), Daguerreia inermis Soares & Soares 1947 in the “Gruta da Lancinha”, Parana, Brazil (Pinto-da-Rocha 1996b), and Pachyloi- dellus goliath Acosta 1993 in “Pampa de Achala”, Cordoba, Argentina (Acosta et al. 1995). In southeastern Brazil, the composition of harvestmen fauna of the Atlantic Rain Forest is different in each mountain chain and more than 50 species can be found in each area of endemism (Pinto-da-Rocha 1999). Most spe- cies are endemic, occupying small areas, nor- mally one mountain chain or some caves (Pin- to-da-Rocha 1999). The cavernicolous harvestmen do not vary in abundance through the year, and their populations seem to be more stable than epigean populations (Gnas- 208 MESTRE & PINTO-DA-ROCH A— BIOLOGY OF HARVESTMAN ILHAIA IN BRAZIL 209 pini 1996; Pinto-da-Rocha 1996a; 1996b), probably because epigean climates are more variable than inside caves. Thus, we expect that epigean harvestmen populations will vary in abundance through the year. The epigean species llhaia cuspidata Roewer 1913 has a wide distribution, occur- ring in southern to southeastern Brazil from the state of Rio de Janeiro to Parana. It in- habits leaf litter of subtropical humid forests and synantropic environments, such as second growth and forest fragments inside cities. In this study we examinate the population dy- namics and intra and interspecfic interactions of this harvestmen species in an urban site, intending to understand possible seasonal var- iations in abundance and other biological as- pects. METHODS Study site (Fig. 1). — This study was car- ried out in an Araucaria forest fragment, lo- cated in the “Museu de Historia Natural do Capao da Imbuia” east of Curitiba county, Brazil (25'25"48°S, 49'16"15°W). This forest fragment comprises 30,000 wA that was iso- lated in the last 50-70 years and today is sur- rounded by houses, prior the park foundation. The Araucaria forests in south Brazil are char- acterized by trees averaging 25-30 m high, with great biomass of large Gymnospermae Araucaria angustifolia (Bertol. 1898.) Kuntze. Open understories are dominated by shrubs and vines, and in secondary forests, the case of this study, the ground is mainly covered by leaves and fallen trees, where harvestmen can live. Subtropical Humid Mesotermic charac- terizes the climate of the region, with a warm season and a cold season with frequent frosts (May-September). The mean temperature in the warm season is below 22 °C and in the coldest months are below 12 °C. The annual mean is 17 °C. Rainfall is between 1300 and 1500 mm per year (105-190 mm each month from September-March and 78-102 mm from April-August) and relative humidity averages 85% (Mack 1981) (Fig. 2). The meteorologi- cal data (Fig. 2) used in this study were ob- tained from the “Estagao Meteorologica do Centro Politecnico” (located 5 km from the study site) supplied by the “Sistema Meteo- rologico do Parana (SIMEPAR)”. Capture-recapture. — Individual harvest- men were collected from June 1 997 to August of 1998 by one person, during daylight, with intervals of 21-30 days between each one of the 21 samples. The time spent in the held taking samples varied from 5-16 h, depending on the total number of captured animals. The study site was arbitrarily divided into 23 sam- pling points of strips of 100 m^ (Fig. 1). At each sample point all harvestmen were care- fully collected (under fallen trunks, boards, bricks, etc.) with forceps and a flashlight for searching in crevices. All adults and last nymphal stages (i.e., harvestmen without ar- olium, see Munoz-Cuevas 1971) encountered were captured. The immature stages of /. cus- pidata Roewer 1913 and other harvestmen species observed were counted, but not cap- tured or marked. After capture, the harvest- men (adults and last nymphal stages) were sexed (differentiated mainly by heavy arma- ture of male leg IV, weak on females), counted and marked with acrylic or plastic ink on the back region of the dorsal scute or on the fem- ora of the fourth leg in speciflc combinations of colors for each sample date. This ink did not harm the individuals and after several re- captures the marks were still easily recogniz- able (see Appendix 1). The marked individu- als were released at the same place where they were collected. Voucher specimens were de- posited in the Museu de Zoologia da Univer- sidade de Sao Paulo and Museu de Historia Natural “Capao da Imbuia”. Shelter preferences. — The 23 sample points were characterized on the basis of the quantity of possible shelters present in each sample site (100 m^) that could be influencing the local abundance of harvestmen. The shel- ters were classified in 4 categories for subse- quent statistical analysis: trunks, stones, bricks and trash (objects of inorganic origin, plastic objects). We compared separately the number of adult harvestmen observed per sample point, in habitats with high (H), medium (M) and low (L) concentration of different shelters by Kruskal- Wallis and Dunn’s Tests. Behavioral observations. — Prior to our population study we collected harvestmen on three dates (2 1 March, 1 1 April and 5 May 1997). At these times captured individuals were individually marked with a small drop of acrylic ink. These data were not considered in the estimates of population size, but they were used for observations of individual movements. During each sample period we re- 210 THE JOURNAL OF ARACHNOLOGY Figure 1. — Map of the study site, Curitiba, Brazil. 50 m 1,2 Sampling points Buildings E Vegetation ■i Water TRAILS corded the number and sex of aggregated in- dividuals. We considered as ''aggregated” harvestmen that had legs in contact with one or more others harvestmen. We established five categories of aggregations: (a) males only, (b) females only, (c) both males and fe- males, (d) males, females and immatures and (e) multi-species aggregations, with individu- als of L cuspidata and other harvestmen spe- cies. We analyzed differences between num- bers of individuals in aggregates by seasons. We made two night observations in the study site (on 20 March and 13 July 1998). We counted the active individuals observed in a 100 m transect (width 5 m), in periods of 15 minutes, from 1700 h-2000 h. Behavioral and foraging observations were made in a hexag- onal terrarium of glass (50 x 40 cm) with ap- proximately 2 cm of soil, three shelters and two plastic receptacles with water. This ter- rarium housed 13 individuals of Ilhaia cus- pidata, one female of Discocyrtus sp.2, one male of the spider Polybetes pitagoricus (Holmberg 1875) (Araneae, Sparassidae), one female of the spider Ctenus sp. (Araneae, Ctenidae) and two diplopods. Also included were crickets, cockroaches, isopods and slugs. Statistical methods. — The population size of /. cuspidata was estimated by Fisher-Ford and Jolly algorithms (Begon 1979). Both methods require several phases of marks and recaptures. In the Fisher-Ford method, the population size is estimated with the assump- tions that the relationship between marked and total individuals found during the sampling period is the same for the whole population and that survival rate is constant and indepen- dent of age. In this study, only the adults and last nymphal stage were sampled. Jollys meth- od considers only the most recent recaptures and the oldest are ignored (Begon 1979). This method calculates a survival rate for each sample (Begon 1979). Although the marks on the subadult stage could be lost, these indi- viduals were also included in the population estimates because it is difficult to separate them from the adults in the field, and because this stage can span more than 9 months in some harvestmen species (Gnaspini 1996). The data obtained using each method were MESTRE & PINTO-DA-ROCHA— BIOLOGY OF HARVESTMAN ILHAIA IN BRAZIL 211 100 n Figure 2. — Mean, maximum and minimum temperature (above) and relative humidity (below) in study site. Autumn began on 20 March, winter on 21 June, spring on 23 September and summer on 21 December. 212 THE JOURNAL OF ARACHNOLOGY 16000 Figure 3. — Estimates of population size of adults and subadults of Ilhaici ciispidata by Jolly and Fisher Ford’s capture-recapture methods in an Araucaria forest fragment in Brazil. compared with a Mann- Whitney U test. Com- parisons of population size between seasons were made with ANOVA and Tukey's q test. The density of harvestmen at the study site was determined by dividing the population es- timate divided by the total area of the frag- ment (30000 m^). We used chi-square to test whether the sex ratio deviated from 1:1 and the association between juveniles and adults observed in the four seasons sampled. We used KruskaL Wallis (KW) and Dunn’s Q to test for possible differences in harvestmen abundance in habitats with distinct concentrations of shelters, seasonal variations in abundance of other species of harvestmen, and seasonal dif- ferences in abundance of aggregations. RESULTS Population dynamics* — The estimated population sizes of Ilhaia cuspidata were sim- ilar for both population estimation methods, Jolly and Fisher-Ford (Maen-Witney U = 178.00 P = 0.624) (Fig. 3, Appendix 1). The Fisher-Ford method presented less variation when compared with the Jolly estimate, prob- ably due to the importance of all capture and recapture data points in the population size es- timates. On the other hand, the Jolly method analyzed data more consistently for the day of captures/recaptures (Begon 1979). Herein, these data will be discussed mainly with the Fisher-Ford estimation method. The population estimates ranged from a low of 1,429 individuals in the second sample (winter, 7 June, 1997), to a maximum of 14,445 in the sixteenth sample (autumn, 28 April, 1998). A slight and progressive popu- lation growth was observed from June to No- vember of 1997, when the estimated values oscillated between 1,429 and 2,661 individu- als (Fig. 4). There were significant seasonal differences in the mean population size of Il- haia cuspidata showed a significant difference in winter and autumn, spring and autumn, and summer and autumn, (ANOVA P < 0.0001, Tukey q = 16.3 for winter vs autumn, 14.8 for spring vs autumn, 11.34 for summer vs autumn, P < 0.001). Population size remained low from the end of the winter (June) to the spring (November) and increased quickly in the following four months, during the spring and the summer (increasing 300%). In sum- mer (January 1998) the population increased very rapidly, and in the period of four months, increased from 2,661 to 14,445 adult/subadult estimated individuals. The highest estimate occurred on the sixteenth sample date (28 April 1998), when 1,738 individuals were captured of which 1,227 were marked (Fig. 3, MESTRE & PINTO-DA-ROCHA— BIOLOGY OF HARVESTMAN ILHAIA IN BRAZIL 213 Table 1 . — Total number of males (M) and females (F) of harvestmen species observed on each sample date at the study site. D.L Discocyrtus spi, D.2. Discocyrtus sp25 Tr. Tricommatidae, G. sp. Geraecor- mobius sp., /x. = Ilhaia cuspidata marked, jv. = juveniles, t = dead observed. Sample date DJ. M D.L F D.2. M D.2. F Tc M + F G. sp. M G. sp. F /.c. M /.c. F I.c. jv. Led 07 June 1997 i 1 0 0 0 1 0 157 135 8 0 28 June 1 1 0 0 1 0 0 113 73 1 0 21 July 1 6 0 0 3 0 0 107 107 5 0 1 1 August 0 2 1 0 4 1 1 131 131 2 0 05 September 3 3 0 0 8 0 0 112 113 5 5 22 September 4 6 1 1 8 2 0 127 159 0 1 13 October 5 8 1 1 3 1 2 92 95 50 1 03 November 8 13 1 1 4 2 2 78 134 144 1 24 November 6 9 0 0 5 0 0 104 96 334 2 14 December 4 5 i 1 6 1 0 149 136 226 1 05 January 1998 8 12 0 3 7 2 0 274 224 154 1 25 January 16 11 0 0 4 4 5 300 267 141 3 24 February 23 22 0 2 18 2 1 482 497 87 1 19 March 13 18 1 2 6 1 i 539 624 94 8 08 April 18 16 1 2 7 1 2 696 763 76 17 28 April 5 9 3 2 11 0 0 567 660 38 20 23 May 6 7 0 0 5 2 1 390 445 30 22 15 June 3 10 1 2 8 0 1 324 374 23 34 13 July 3 4 1 0 7 3 2 184 206 25 37 03 August 4 8 0 0 13 2 2 151 159 24 34 04 September 4 7 1 0 8 1 0 92 128 13 72 Total 136 178 13 17 136 26 20 5169 5526 1480 259 Appendix 1). The rates of recapture of marked individuals varied from 16% (28 June 1997) to 41% (24 November 1997) of the total cap- tured in daylight. The density of Ilhaia cuspidata in the study site varied between 0.05 adults/m^ (in the win- ter) to 0.47 adults/m^ (in the summer). The adult sex ratio was 1:1 for the entire study period (A^ - 33.82 df = 20 F < 0.027). How- ever, when the seasons were analyzed sepa- rately, only in spring the was sex ratio 1 : 1 (JG = 9.37 df = 3 P < 0.024), in other seasons these data showed alternating sex predomi- nance (winter I: males > females = 5.13, df = 4, F < 0.27, summer: males > females X ” 6.99 df = 4 F < 0.14, autumn: males 0.05). Aggregations that contained from 6-9 individuals show a signihcant increase in summer (Su) and au- tumn (Au) when compared with spring (Sp) (KW = 16.77, Dunn Qs„s, = -14.87 P < 0.01, = -13.37, P < 0.05). We also ob- served a signihcant increase in the number of aggregations with more those 10 individuals when comparing spring and autumn (KW = 15. 0005^, = —15.00, P < 0.05). The analysis of all aggregations together shows similar re- suits (KW = 14.72, Qs„s„ = -13.62, = -14.12, P < 0.05). Thus, aggregations larger than six individuals were more abundant in summer and autumn. Individual movements and life span. — During the three previous collections, taken before the sampling period, 238 individuals of /. cuspidata were individually marked (num- bered). We recaptured 50 of those specimens during the 335 sample days. About 50% of these individuals were found in the same place that they were marked, 46% were found close to these sites (between 10 and 20 m) and the remained 4% were found approximately with- in 60 meters from the point that they were captured. These data show very low vagility of this opilionid species. The number of har- vestmen recaptured decreased constantly across the study period and the last recaptures were made just one year after they had been marked, in the 15'*’ sampling (April 1998). Longevity of adults is more than 455 days. Natural history. — Field observations showed that /. cuspidata is practically motion- less during the day, taking shelter in the in- terior or under fallen trunks, stones and pieces of wood. Individuals were also observed tak- ing shelter under plastic objects and other trash. During the two nocturnal samples, one in the warm season (March 1998), and the other in the cold season (June 1998), we ob- served that the harvestmen began activity ear- ly in the night, about 15 minutes after sunset. An increase in the number of active individ- uals was observed 30 minutes after sunset, when it was almost dark (light intensity was 02 lux). On 15 June 1998, just after sunset, from 1730-1745 h, no active harvestmen were observed. The observations from 1750- 1905 h revealed an incremental increase in harvestmen activity. From 1750-1805 h, all the individuals (16) of /. cuspidata seen were active; 18 were observed from 1810-1825 h; 38 from 1830-1845; and 44 from 1850-1905 MESTRE & PINTO-DA-ROCHA— BIOLOGY OF HARVESTMAN ILHAIA IN BRAZIL 215 Table 2. — Number of aggregations of Ilhaia cuspidata composed of only males (M); only females (F); males, females and juveniles (M + F T jv.); and interspecific aggregations (intrsp.). Date M F M + F M + F T jv. Intrsp. Total 07 June 1997 4 6 19 1 1 31 28 June 2 1 9 0 3 15 21 July 2 3 15 0 1 21 1 1 August 1 0 7 1 1 10 05 September 3 4 17 0 4 28 22 September 0 3 7 0 0 10 13 October 0 1 9 2 4 16 03 November 0 0 7 1 2 10 24 November 0 0 3 1 1 5 15 December 2 1 6 20 2 31 05 January 1998 2 3 12 10 5 32 25 January 2 3 16 12 10 43 24 February 1998 9 5 36 12 8 70 19 March 1998 3 7 46 10 5 71 08 April 1998 2 5 37 12 3 59 28 April 1998 1 2 31 3 5 42 21 May 1998 1 3 20 2 2 28 Total 34 47 297 87 57 522 % 6.50 9.20 56.90 16.50 10.90 100.00 h. Activity during daylight was never ob“ served outside the shelters. In the field, individuals of /. cuspidata were observed feeding on small adult lepidopter- ans, bird and rodeetai feces (probably Crice- tidae). In captivity, individuals accepted sev- eral kinds of vegetables and fruits (e.g. papaya, banana, peach, beet, and carrot). We observed animals coexisting with L cuspidata, such as toads, worms (Oligochaeta and Hirudieea), crickets, spiders, pulmonate moilusks, pseudoscorpions, coleopteraes (Pas= salidae, Staphylinidae) and other insects. Fre- quently, we observed the presence of small crickets (Phalangopsidae) and a species of pulmonate mollusk near /. cuspidata. The spider Scytodes sp. (Araneae, Scytodi- dae), was observed preying on a young L cus~ pidata, and feeding on the remains of an adult exoskeleton. One dead individual of L cuspi- data was found in the web of a Theridiidae spider. Besides these predators, it is possible that toads {Bufo sp.), ants {Pachycondyla sp. and Odontomachus sp.) and some spiders (e.g. Ctenus sp.) in the study site could prey on harvestmen. Egg-batches of L cuspidata were rarely found (even previous to the peak of young recruitment), probably due to the difficulty of locating them among the leaf litter. The eggs measured approximately 1mm in diameter, were white in color, opaque and sticky. They were laid under fallen leaves in clusters of up to 20 and did not receive any additional care by the female. An egg batch containing 8 eggs was collected in the field and maintained in a plastic box with leaves and wet cotton in the bottom. After 3 weeks, four nymphs hatched. Other harvestmen species. — Other har- vestmen species observed coexisting with Il- haia cuspidata were two species of Discocyr- tus (Gonyleptidae, Pachylinae, one black, sp. 1 and another red, sp.2), one species of Gerae- cormobius sp. (Gonyleptidae, Gonyleptinae), and one species of an unidentified genus of Tricommatidae. We observed significantly higher abundance of Discocyrtus sp.l in sum- mer (Su) and autumn (Au) when compared with the first winter studied (Wi) (KW = 13.93, P = 0.007; QwiSu ~ “ 12.4, Qwiau ~ ~~ 13.2, P < 0.05). We also observed an increase in Tricommatidae (24 February 1998), Ger- aecormobius sp. (25 January 1998) and Dis- cocyrtus sp.2 (28 April 1998) (Fig. 5 & Table 1). These species seem to have a population increase in the same season as Ilhaia cuspi- data and Discocyrtus sp.l, however there is 216 THE JOURNAL OF ARACHNOLOGY 50 , -Discocyrtus sp1 ■ Discocyrtus sp2 Thcommatidae - -x- - Geraecormobius sp. Figure 5. — Total number of four species of opilionids (adults and immatures) observed in an Araucaria forest fragment in Brazil. insufficient data to strongly support this ob- servation. DISCUSSION Biological activities of populations, mainly in tropical regions, can be linked to seasonal parameters such as temperature and rainfall (Opler et al. 1976; Winemiller 1989; Machado & Oliveira 1998). The harvestman Cynor- toides cubanus Banks 1909 (Cosmetidae) de- crease post-embryonic development at higher temperatures (Juberthie 1972). In the same way, there could be a specific temperature that favors Ilhaia cuspidata juveniles to survive in higher numbers until the adult stage. Our data show that the high number of juveniles ob- served in spring and summer was associated with the high number of adults observed in summer and autumn, suggesting that there is a definite period of successful reproduction, with a subsequent adult population increase. The constant presence of juveniles during the whole year, suggests a continuous repro- duction with a peak of emergence just after winter or, delayed hatching. Young and adults often are under different selection pressures. and reproductive effort reflects the environ- mental factors that put pressure on the adults as well as on the juveniles (Pianka 1994). In this respect, a species with high mortality needs a con*esponding high fecundity, persist- ing despite the great decline of the population (Pianka 1994). Very low temperatures should eliminate a great part of the population, main- ly the juveniles. However, there could be a possible decrease in activity and/or movement to other shelters (under soil surface or fallen leaves, available all year) during the coldest times, resulting sub- or super-estimation in certain seasons. In contrast to the cavernicolous harvest- men, such as Goniosoma spelaeum (studied by Gnaspini 1996), Pachylospeleus strinatU (studied by Pinto-da-Rocha 1996a), and Da- guerreia inermis (studied by Pinto-da-Rocha 1 996b), the epigean I. cuspidata had great sea- sonal variation in population size. In caves the temperature is less variable than epigean hab- itats (Culver 1982) and probably the seasonal temperature variation observed in the Arau- caria forest influences this population. The al- MESTRE & PINTO-DA-ROCHA— BIOLOGY OF HARVESTMAN ILHAIA IN BRAZIL 217 ternating sex ratio of the seasons could reflect these fluctuations. The significant difference in harvestmen abundance in habitats with varying concentra- tions of trunks and bricks indicates that these shelter preferences can be linked to microhab- itat conditions promoted by these materials. Wood and clay provide shelters with less tem- perature variation than stones and plastic ob- jects. The great abundance of this harvestmen species in this study site could be linked to their adaptation or preference for these abun- dant shelters. Capocasale & Bruno-Trezza (1964) ob- served that behavior of harvestmen was relat- ed to changes in the temperature. They dem- onstrated that the number of aggregated individuals of Acanthopachylus aculeatus (Kirby 1818) (Gonyleptidae) was inversely proportional to the temperature, with the larg- est aggregations occurring in lower tempera- tures. We observed that /. cuspidata increase in number of larger aggregations during the summer and autumn compared with the spring, thus the number of aggregated individ- uals was not increasing with lower tempera- tures. So, results suggest that the increase of aggregated individuals is not influenced by temperature, but probably by the population density. Aggregation occurs at the limited number of favorable shelter sites(with no light and high moisture, like bricks). In contrast to Goniossoma longipes (Ma- chado & Oliveira 1998) and Goniosoma prox- imum (Ramirez & Giaretta 1994), which guard their egg batches (60-210 eggs) or ju- veniles, parental care of the low number of egg batches (approximately 20 eggs) was not observed for Ilhaia cuspidata. We can not say if /. cuspidata only lays about 20 eggs during the entire reproductive period or lays several egg batches in different shelters as a way to avoid predation. An increase in active individuals at dusk demonstrates that /. cuspidata is nocturnal, forages near shelters, and are restricted to small areas (around 50 m^). This observation supports the concept of a small home range for gonyleptideans, as was also observed for Goniosoma spelaeum, Daguerreia inermis and Pachylospeleus strinatii (Gnaspini 1996; Pinto-da-Rocha 1996a, 1996b). These harvestmen have a very broad diet, as was also observed for Acanthopachylus aculeatus (Capocasale & Bruno-Trezza 1964), Pachyloidellus goliath (Acosta 1995), Pachy- lospeleus strinatii (Pinto-da-Rocha 1996) and Goniosoma spelaeum (Gnaspini 1996), G. longipes (Machado & Oliveira 1998), and Da- guerreia inermis (Pinto-da-Rocha 1996b). Be- sides small live animals and plants, we ob- served this species feeding on the feces of birds and rats, an observation also document- ed by Hillyard & Sankey (1989) for another species, thus reaffirming an omnivorous diet for harvestmen. The /. cuspidata population in this study experienced high fluctuations in the number of individuals, a probable delayed response to climate fluctuations. These population fluctu- ations were also observed for other harvest- men species in the study site, and possibly these harvestmen have similar responses to these environmental conditions. ACKNOWLEDGMENTS We are grateful to Anderson K. Eujioka, Antonio Serbena, Luis Coltro Jr. and Renata C. C. Sousa for help with fieldwork, and to Glauco Machado for suggestions on the man- uscript. We thank Jose Roberto Varani for help with statistical analyses. We are grateful to the director of the “Museu de Historia Nat- ural Capao da Imbuia” for permitting access to the study site and to Pedro Gnaspini who kindly permitted the use of his computer mark-recapture program. LITERATURE CITED Acosta, L.E., E. Pereyra & R.A. Pizzi. 1995. 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Description of male of Daguerreia inermis Soares & Soares, with bio- logical notes on population size in Gruta da Lan- cinha, Parana, Brazil (Arachnida, Opiliones, Gonyleptidae). Revista Brasileira de Zoologia 13(4):833-842. Pinto-da-Rocha, R. 1999. Opiliones. In C.A. Joly, and C.E.M. Bicudo (org.). Biodiversidade do es- tado de Sao Paulo, Brasil: smtese do conheci- mento ao final do seculo XX. Volume V. Inver- tebrados terrestres. FAPESP. Ramires, E.N., and A. A. Giaretta. 1994. Maternal care in a neotropical harvestmen, Acutisoma proximiim (Opiliones, Gonyleptidae). Journal of Arachnology 22:179-180. Savory, T.H. 1938. Notes on the biology of har- vestmen. Journal of Queckett Microscopical Club. 1:89-94. Winemiller, K.O. 1989. Patterns of variation in the life history among South American fishes in sea- sonal environments. Oecologia 81:225-241. Manuscript received 11 November 2002, revised 29 July 2003. MESTRE & PINTO-DA-ROCHA— BIOLOGY OF HARVESTMAN ILHAIA IN BRAZIL 219 Appendix 1. — Number of captured (CAP), released (REE) and recaptured (D1 = day, D2 = 22, second day of recapture, D3 = 45, third day of recaptures . . .) adults and subadults of Ilhaia cuspidata in Curitiba, Parana, Brazil. Recaptures Day CAP REL D1 D2 D3 D4 D5 D6 D7 D8 D9 1 292 292 22 222 222 36 45 272 272 23 35 66 338 338 24 16 36 91 297 295 17 9 26 30 108 380 380 8 10 17 19 40 129 298 290 6 7 12 15 30 33 150 332 327 6 4 11 17 22 28 27 171 339 339 7 7 13 13 18 26 15 40 191 396 395 2 5 9 3 13 9 12 31 26 213 671 670 3 2 9 8 16 10 13 23 20 232 850 838 5 0 7 2 9 14 10 24 22 263 1314 1297 4 4 6 6 12 11 9 22 21 286 1526 1526 4 4 6 7 9 6 8 19 18 306 1896 1896 2 2 7 4 8 5 9 11 9 326 1738 1736 4 2 8 7 5 13 3 6 1 1 349 1304 1304 5 3 3 4 7 8 6 4 6 373 1105 1105 4 1 6 3 6 6 1 6 5 402 715 714 1 0 2 2 2 2 4 4 3 423 562 562 0 0 2 0 1 2 2 3 3 455 403 403 2 1 0 0 1 1 3 2 2 DIO 68 35 31 22 3 7 3 4 4 2 1 THE JOURNAL OF ARACHNOLOGY Appendix 1. — Extended. Recaptures Dll D12 D13 D14 D15 D16 D17 D18 D19 D20 143 89 103 42 68 150 30 50 108 189 21 24 57 114 227 17 22 37 59 120 165 7 8 32 41 83 92 102 8 9 15 32 44 64 52 77 2 4 10 27 27 37 30 50 5 4 3 18 16 22 15 33 50 28 26 2004. The Journal of Arachnology 32:221-230 PHENOLOGY OF LINYPHIIDS IN AN OLD^GROWTH DECIDUOUS FOREST IN CENTRAL ALBERTA, CANADA Christopher M, Buddie: Department of Natural Resource Sciences, McGill University, Macdonald Campus, 21,111 Lakeshore Road, Ste. Anne de Bellevue, QC H9X 3V9 Canada. E-mail: chris.buddle@mcgilLca Michael L. Draney: Department of Natural and Applied Sciences and Cofrin Center for Biodiversity, University of Wisconsin-Green Bay, Green Bay, Wisconsin 54311 USA ABSTRACT, Spiders in the family Linyphiidae are numerically dominant and show remarkably high diversity in northern forests, but relatively little is known about their phenology in northern latitudes of North America. We report a phenological summary of close to 6,000 individual linyphiids representing 17 species. These were collected by pitfall trapping during two snow-free seasons in an old-growth de- ciduous boreal forest in central Alberta, Canada. Three species of approximately the same body size, Allomengea dentisetis (Griibe 1861), Bathyphantes pallidus (Banks 1892), and Lepthyphantes intricatus (Emerton 1911), dominated the sample, and showed three distinct patterns of peak activity. This suggests temporal stratification as a possible mechanism that explains their co-existence. Four less commonly collected species within the same genus (Walckenaeria) showed similar seasonal segregation in periods of peak activity. Comparisons with other literature suggest the general phenology of many linyphiids is conserved across continental and global scales. Keywords: Species co-existence, life-history, boreal forest, pitfall trapping The family Linyphiidae (sensu lato) is the second most diverse spider family globally (Coddingtoo & Levi 1991), and is notable in attaining greatest diversity in north temperate latitudes rather than towards the equator. As such, linyphiids dominate northern spider fau- nas (Helsdingen 1983). A notable biological feature associated with this high northern di- versity is a flexibility of phenology (seasonal sequence of life history events) among mem- bers of the family. Lieyphiid species display most of the phenological patterns known among spiders: multivoltine and ueivoltine strategies predominate in warmer climates, and the reproductive period of iinivoltines may occur in various seasons and for varying durations (Merrett 1969; Berry 1971; Draney 1997a, b; Draney & Crossley 1999). North- ward, biennial and mixed annual-biennial strategies (in which some members of a pop- ulation are annual and some are biennial) be- come more common (Schaefer 1977; Toft 1976, 1978). Different species overwinter as juveniles, adults, or (more rarely), eggs (Schaefer 1976, 1977), and some species ac- tively feed and even reproduce during the northern winter (Huhta & Viramo 1979; Aitchison 1978, 1984). Phenological flexibility has also been doc- umented within lieyphiid species, with longer cycles being displayed by more poleward pop- ulations (Almquist 1969; Toft 1976). Besides being a response to climate, phenological pat- terns are also associated with other ecological factors including vertical stratification of the spider (Toft 1978) and habitat type (Draney & Crossley 1999). Certainly, within any climatic regime, a variety of phenological patterns may be displayed by different linyphiid species. Most of the work on linyphiid phenology has examined western and northern European populations experiencing relatively mild mar- itime-iefluenced climates (Juberthie 1954; Tretzel 1954; Almquist 1969; Merrett 1969; Toft 1976, 1978; De Keer & Maelfait 1987, 1988; Hauge 2000). With the exception of work in Finland (e.g., Huhta 1965, 1971; Palmgren 1972; Memela et al. 1994), less is known about linyphiid phenological responses to harsher continental boreal climates, such as 221 222 THE JOURNAL OF ARACHNOLOGY our study site in central Alberta, where sea- sonal variation is extreme. Some work has been done on the subnivean winter activity of spiders in these continental climates (Aitchi- son 1978, 1984). The present work examines growing season phenology of linyphiids in a sub-boreal climax deciduous forest. Our ob- jectives include describing the phenological patterns of the common linyphiids as well as attempting to identify factors that might ex- plain the variation in phenological patterns among the species in this system. We ac- knowledge that conclusions about phenology may be limited to our study location, sample period, and study years, but nonetheless hope to increase knowledge about a poorly under- stood component of the boreal spider fauna. METHODS Study site, — The study forest is located at the George Lake Field Station, located 75 km northwest of Edmonton, Alberta, Canada (53°57'N, I14°06'W). This old-growth decid- uous forest has been left relatively undis- turbed for over 100 years. The work described here was part of two larger projects investi- gating the relationship between ground-dwell- ing spiders and fallen logs, or downed woody material (Buddie 2001 a, b). The first project occupied an area of approximately 2.5 ha at the northeast section of the field station. The second project comprised about 3.5 ha in the northwest portion of the forest. Both projects were conducted in homogenous regions of this boreal mixed-wood forest, which is dominated by two Populus species: trembling aspen (Populus tremuloides Michx.) and balsam poplar {Populus bahamifera L,). Less com- mon tree species in the area include birches {Betula papyrifera Marsh, and B, neoalaskana (Sarg.)), white spruce (Picea glauca (Moench)) and black spruce {P. mariana (Mill.) BSP). Further vegetation details can be found in Niemela et al. (1992). Sampling and spider identifications. — Ground-dwelling spiders were sampled with 88 pitfall traps in 1998 and 232 pitfall traps in 1999. All 88 traps in 1998, and 88 of the pitfall traps in 1999 were white circular (11 cm diameter) plastic containers sunk into the ground with the lip flush to the substrate sur- face (Spence & Niemela 1994); 2-3 cm of preservative was used in the traps (silicate- free ethylene glycol) and a plywood roof mea- suring 15 X 15 cm was elevated 2-3 cm above the traps to prevent flooding and trap disturbance. The additional 144 pitfall traps used in 1999 were smaller (6 cm diameter), made of clear plastic, and were covered with a circular plastic roof (11 cm diameter). Larg- er traps were placed 10-15 m apart; smaller pitfall traps were all 2 m from an adjacent trap (see Buddie (2001 a, b) for complete details regarding sampling design). There has been some debate over the use of pitfall traps to sample ground-dwelling ar- thropods, as such traps are biased by the ac- tivity of the organism, and are influenced by such factors as trap material, type of preser- vative and color (e.g.. Luff 1975; Adis 1979; Curtis 1980; Merrett & Snazell 1983; Topping 1993), However, pitfall traps have been shown to efficiently sample ground-dwelling spiders (e.g., Uetz & Unzicker 1976; Draney 1997 a, b; Buddie et al. 2000), and provided that no statements are made about absolute density, pitfall traps can be used to quantitatively as- sess periods of peak activities of male and fe- male spiders (Toft 1976, 1978; De Keer & Maelfait 1987; Draney & Crossley 1999). Sampling was continuous over the snow- free season in both years; in 1998 pitfall traps were placed in the forest on 4 May and re- moved on 10 September. Pitfall traps were opened from 20 April until 24 September in 1999. This provided 126 days of continual pit- fall trapping in 1998 and 154 continuous trap- ping days in 1999. There were eight collection times (approximately every 15-20 days) in each year. Samples were sorted and stored in 70% eth- anol. All adult spiders were identified to spe- cies with nomenclature following Platnick (2003), and Buckle et al. (2001) for liny- phiids. Voucher specimens of all taxa have been deposited in the Strickland Entomologi- cal Museum (Department of Biological Sci- ences, University of Alberta) and the Northern Forestry Centre Arthropod Collection, both in Edmonton, Alberta, Canada, Data standardization and analysis. — In- creasing sampling effort (i.e., number of traps) corresponds to an increase in the total catch of individuals (Niemela et al. 1986). It was therefore necessary to standardize the collec- tion data from 1998 and 1999 to account for variation in trapping effort. Without this stan- dardization, it would be difficult to separate BUDDLE & DRANEY— LINYPHIID PHENOLOGY 223 true differences in relative abundance from simply differences in sampling effort. Prior to data standardization, however, species by sample accumulation curves were constructed using Estimates Version 6.0 (Colwell 1997). This was to ensure that linyphiid species rich- ness from 1998 (88 pitfall traps) and 1999 (232 traps) had leveled off, and thus repre- sented complete samples. Only in this case would further standardization be justified. When this criterion was met, and when col- lections from different years were compared, pitfall trap data were adjusted to make catch data relative to a predetermined number of pit- fall traps; since there were 88 traps in 1998 and 232 traps in 1999, data for both years were standardized to catches per 160 traps (average number of traps per year). Samples in 1998 were multiplied by 1.82 (i.e., 160/88), and samples in 1999 were multiplied by 0.69 (i.e., 160/232). Data were not standardized to account for differences in pitfall trap size (i.e., 6 cm versus 1 1 cm diameter) as Work et al. (2002) show catches of spiders do not vary significantly between these trap sizes. Linyphiid species represented by fewer than 15 individuals in the collection were excluded prior to analyses as these would be too few to adequately assess phenological patterns. It was assumed that periods of peak male and female activity correspond to the peak repro- ductive period for the species (e.g., De Keer & Maelfait 1987; Draney 1997 a, b). Graph- ical analysis was used to assess this reproduc- tive period and to evaluate the year-to-year variation in catches of the common ground- dwelling spiders. Results were compared to other published records and to information available on the same or closely related spe- cies from different geographic regions. Addi- tionally, we measured the carapace width (CW) of our study spiders to ascertain overall spider size (Hagstrum 1971) to determine if any phenological patterns differed by species size. For these measures, a sub-set (i.e., 3-5 individuals) of both males and females were measured under a dissecting microscope fitted with an ocular micrometer. These measures were used purely in a relative sense, and thus sample sizes were small, and measures of var- iance were not used. RESULTS A total of 5,944 individuals representing 50 species of linyphiids were collected. Liny- phiids represented 32% of the total number of spiders collected experiment-wide, and 46% of the total number of species collected (see Buddie 2001a for complete species list). Of the linyphiids, 17 species were represented by > 15 individuals, and these accounted for > 98% of the total number of linyphiids col- lected (Table 1). Three species accounted for most of the linyphiids (87.2% of the total lin- yphiids collected): Allomengea dentisetis (Griibe 1861), Bathyphantes palUdus (Banks 1892), and Lepthyphantes intricatus (Emerton 1911) (Table 1). Species accumulation curves show that the observed species richness of the linyphiids had leveled off at about 80 samples in 1998 and about 65 samples in 1999 (Fig. 1). This illustrates that we adequately sampled liny- phiids in this study, and comparisons across years are justified, as is standardization to ac- count for sampling effort. On a per-trap basis, about the same number of linyphiids were collected in 1998 (mean (± SE) of 21.53 ± 1.39 spiders per trap, n = 88)) as in 1999 (17.02 ± 0.57, n = 232). Stan- dardized to 160 traps, we collected 3445.5 lin- yphiids in 1998 and 2722.8 in 1999. When the total male and female catch of the 1 7 linyphiid species is plotted by year and sampling time, it is apparent that males were more common in pitfall traps than were females. The peaks in female activity generally corresponded with the peak in male activity (Fig. 2). In both years female linyphiids were most commonly collected late in the season; males were more common in pitfall traps late in the season in 1998, but were less variable across collection dates in 1999 (Fig. 2). Collections of the three common linyphiids showed three different phenological patterns, and these patterns were similar for both col- lection years (Fig. 3), Allomengea dentisetis was most active in August and September, B. pallidus in late July-August, L. intricatus in June. In general, females were less frequently collected than males, but their peak in activity corresponded closely to male activity (Fig. 3). When the phenologies of all 17 linyphiid species are depicted, it is clear that periods of peak activity vary both in when males and females occur, and in the length of time both males and females are active (Fig. 4). For ex- ample, males and females of L. intricatus, B. pallidus, and Microneta viaria (Blackwall 224 THE JOURNAL OF ARACHNOLOGY Table 1. — Mean carapace width (CW, mm), number of females, and number of males, for 17 species of Linyphiidae collected by pitfall traps in 1998 and 1999, in a deciduous forest in north-central Alberta. Species arranged by size (descending). Species CW Females Males Total Pityohyphantes costatus (Hentz 1850) 2.80 19 20 39 Allomengea dentisetis (Griibe 1861) 1.36 1068 1719 2787 Neriene clathratci (Sundevall 1830) 1.28 2 30 32 Helophora insignis (Blackwall 1841) 1.20 41 30 71 Lepthyphantes iutricatus (Emerton 1911) 1.14 124 532 656 Oreonetides vaginatiis (Thorell 1 872) 1.13 13 60 73 Centromems sylvaticus (Blackwall 1841) 1.02 1 14 15 Bathyphantes paJlidus (Banks 1892) 0.88 433 1311 1744 Walckenaeha prominens Millidge 1983 0.82 3 64 67 Walckenaeria castanea (Emerton 1882) 0.81 26 0 26 Microneta viaria (Blackwall 1841) 0.78 12 130 142 Lepthyphantes zebra (Emerton 1882) 0.78 2 30 32 Walckenaeria directa (O. R-Cambridge 1874) 0.77 5 34 39 Sciastes trimcatus (Emerton 1882) 0.66 0 36 36 Walckenaeria atrotibialis (O. P.-Cambridge 1878) 0.66 0 25 25 Pocadicneniis americana Millidge 1976 0.56 0 21 21 Diplocentria bidentata (Emerton 1882) 0.53 2 36 38 Total 1751 4092 5843 Figure 1. — Observed species accumulation curve of species richness by samples (pitfall traps) in 1998 (88 traps) and 1999 (232 traps). Data were re-sampled (randomly, without replacement) 50 times, enw bars are ± 1 SD. BUDDLE & DRANEY— LINYPHIID PHENOLOGY 225 1000- 100-i 10- 1998 Female Male 1999 ■1000 100 10 May June July Aug. Sept May June July Aug. Sept Figure 2. — Total number of male and female linyphiids (17 species) collected by pitfall traps in an old- growth deciduous forest. Note log-scale on axis. Number of individuals standardized to 160 pitfall traps. 1841) m^ere caught in pitfall traps for much of the collection period, whereas many of the smallest linyphiid species collected show re- duced periods of activity early in the season (e.g., Diplocentria bidentata (Emerton 1882), Walckenaeria directa (O.P.-Cambridge 1874) and Lepthyphantes zebra (Emerton 1882)) (Fig. 4). Smaller-bodied linyphiids were sel- dom collected past mid-summer whereas some of the larger-bodied species show higher catches in pitfall traps late in the season (e.g., A. dentisetis, Helophora insignis (Blackwall 1841) and Centrornerus sylvaticus (Blackwall 1841)) (Fig. 4). Species within the same genus also show some differences in periods of peak activity. For example, collections of the four Walckenaeria species never overlap in time, and L. zebra occurs early in the season where- as L. intricatus shows a peak in activity about 3 weeks later (Fig. 4). It should be noted, however, that many of the species depicted in Fig. 4 were relatively rare in the collection (Table 1), so statements about their phenology should be interpreted with caution. DISCUSSION Linyphiids are dominant on the forest-floor in our old-gromTh study forest, in terms of both diversity and relative abundance; they represented almost Tialf the total species col- lected and one third of the number of individ- uals collected. Our work represents one of the few detailed accounts of linyphiid phenology from northern regions of North America. We have presented data that shows three species, of approximately the same relative body size, are remarkably common on the forest floor of north-central Alberta: A. dentisetis, L. intrb catus, and B. pallidus. These species have also been shown to dominate the fauna of boreal- mixed wood forests throughout north-central Alberta (Buddie et al. 2000; Buddie 2001a). An important question is how these linyphiids might coexist on the forest floor, and the phe- nological summary may provide some clues. It has long been suggested that seasonal segregation of similar-sized spiders might pro- mote species co-existence (e.g., Breymeyer 1966; Williams 1962; Uetz 1977), In our work A. dentisetis shows a period of peak activity late in the season (AugusUSeptember), L. in- tricatus early in the season (June), and B. pal- lidus in mid-summer. Thus, temporal segre- gation may be the mechanism that promotes co-existence of these linyphiids in northern boreal forests. We also see this general pattern within the genus Walckenaeria and Lepthy- phantes from our collections. Future work will have to test this hypothesis, and it is difficult to claim generality from our limited collection time, and limited study area. There are clearly other linyphiids that also co-occur with A. dentisetis, B. pallidus, and L. intricatus, but these may not directly interact with the three dominant species as they are either relatively rare in our collections, or are of a smaller body size (e,g., Walkenaeria spe- cies, M. viaria) (Table 1; Fig. 4), or they use herbaceous vegetation as additional foraging sites. For example, H. insignis is commonly collected by sweeping the vegetation in boreal 226 THE JOURNAL OF ARACHNOLOGY Bathyphantes paliidus "1998 May ‘ June ' July ' Aug. * Sept Figure 3. — Total number of male and female Allomengea dentisetis (top), Bathyphantes paliidus (mid- dle), and Lepthyphantes intricatus (bottom) collected by pitfall traps in an old-growth deciduous forest. Note log-scale on axis of top graph. Number of individuals standardized to 160 pitfall traps. mixed-wood forests (Buddie et ai. 2000), and H. insignis and Pityohyphantes costatus (Hentz 1850) have often been observed in webs located in the herbaceous vegetation at our study forest (C.M. Buddie, pers. obs.). These species do use the forest floor as evi- dent from our pitfall trap collections, but their main foraging location may be in the herba- BUDDLE & DRANEY— LINYPHIID PHENOLOGY 227 Pityohyphantes costatus AHomengea dentisetis Neriene ciathrata Helophora insignis Lepthyphantes intricatus Oreonetides vaginatus Centromerus sylvaticus Bathyphantes pailidus Walckenaeria prominens Walckenaeria castanea Microneta viaria Lepthyphantes zebra Walckenaeria directa Sciastes truncatus Walckenaeria atrotibialis Pocadicnemis americana Dipiocentria bidentata #=& * 1} f=0= a- w #4 f}' -B Size May June July Aug. Sept. Figure 4. — Phenological summary of 17 linyphiid species collected over two years (pooled). Horizontal line indicates continuous periods when males (solid) and females (dashed) were collected. Vertical blocks represent peak activity of males (solid) and females (open), represented by weighted average of males or females collected by sampling period. Species size (smallest to largest, moving vertically) was determined by averaging carapace width for males and females. ceous layer. Therefore, other larger-bodied species may interact with the three dominant species, but the frequency of interactions may be relatively low due to vertical habitat strat- ification (e.g., Turnbull 1960; Luczak 1966). We can compare our phenological summa- ries with other published accounts of seasonal activity of linyphiids to determine whether any species or genera show consistent patterns across larger scales. We focus first on research by Niemela et al. (1994) in Finland, as this work was also done in a mature forest in a climate with similar seasonal extremes as found in northern Alberta. Additionally, Nie- mela et al. (1994) rely on pitfall trap data to ascertain peaks in spider activity, making comparisons with our study relevant, and sev- eral of the same species and genera are com- mon to both studies. Many of our results confirm research by Niemela et al. (1994): D. bidentata occurs most commonly in early season, M. viaria is most frequently collected from June-early August, Oreonetides vaginatus (Thorell 1872) is most abundant early in the season, and H. insignis does not appear in collections until August. Therefore, the seasonal occurrence of some linyphiids is conserved, even on differ- ent continents. Centromerus sylvaticus is known to be ac- tive in the winter under the snow layer in cen- tral Canada (Aitchison 1978). The entire ge- nus is apparently winter active, with a cold season reproductive peak (Kronestedt 1968; Merrett 1969; Huhta & Viramo 1979; Draney 1997b). Centromerus sylvaticus has a low op- timal temperature for postembryonic growth, which results in slow growth during the sum- mer months, delaying maturity until late fall or winter (Schaefer 1977). In southern Eng- land, males and females peak in December and January, and females survive until July. Our data, showing high numbers of individ- uals in fall and none in the spring, may indi- 228 THE JOURNAL OF ARACHNOLOGY cate that in harsh winter climates, reproduc- tion occurs before winter, and adults do not generally survive to the next spring. The four species of Walckenaeria in our data set all displayed a stenochronous pattern of adult activity, with short peaks occurring in late winter (Fig. 4, W. directa), June-Juiy (W. prominens), Mid-July (W. atrotibialis), and early August (W. castanea). Examined species within the large genus Walckenaeria all seem to be univoltine (Tretzel 1954; Merrett 1969; Huhta & Viramo 1979; Draney 1997a); whether this trait is constant within the taxon remains to be seen. Several authors (Schmoller 1970; Muma 1973; Doane & Dondale 1979) have suggest- ed that the male pitfall catch peak is the best indicator of the mating period of a species, since males are trapped as a result of their mate-searching behavior. Female catch is re- lated to either foraging behavior in order to obtain food for egg production, or behavior related to oviposition activity. In many species (such as our R costatus, L. intricatus, and M. viaria. Fig. 4) the male peak occurs well be- fore the female peak, although in other species (A. dentisetis, H. insignis, C. sylvaticus, B. pallidus, L. zebra. Fig. 4) the peaks are essen- tially simultaneous. In no species is the female peak well before the male peak; cases with earlier female peaks occur only in species with few trapped individuals, and the pattern could be a result of sampling error (e.g., W, directa, n = 39 and D. bidentata, n = 38, Table 1). Additionally, we found that more males than females were collected in our pit- fall traps. This is also largely attributable to the differential locomotory activity associated with reproduction; males tend to wander ex- tensively in search of mates. In two cases we collected more females of a species than males (Table 1). This may be due in part to oviposition behavior. For example, Toft (1978) suggested that many species, including //. insignis, lay their eggs in the leaf litter, even if they forage higher in the vegetation. Koponen (1987) also reported highly female- biased pitfall catches of two linyphiid species, Hybauchenidium gibbosum (S0rensen 1898) (95% female, n — 127) and Zornella cultri- gera (L. Koch 1879) (100% female, n = 36). Linyphiid spiders are important predators in northern forests, given their ubiquity, abun- dance, and high diversity. Phenological data can be useful to predict times during which species are likely to occur and also allow us to better predict potential biological interac- tions and population responses to human- caused and natural ecosystem alterations, de- pending on the timing of these events. Phenological data is useful but is not known with precision for most linyphiid species. Use- ful phenological insights can be garnered from ecological data collected for other purposes, and we hope future work will further test pat- terns uncovered in our study forest. ACKNOWLEDGMENTS Financial assistance was in part provided by the Natural Science and Engineering Research Council of Canada (NSERC) in the form of a postgraduate scholarship to CMB and a re- search grant to John R. Spence (Dept, of Bi- ological Sciences, University of Alberta). Ad- ditionally, Natural Resources Canada (Northern Forestry Centre) and the Depart- ment of Biological Sciences (University of Alberta) provided indirect support. We also thank Don Buckle for verifying many of the species determinations, and J.R. Spence, D. W. Langor, K. Cryer, G. Pohl, D. Williams, J. Jacobs, and A. Graham assisted with various aspects of this work. LITERATURE CITED Adis, J. 1979. Problems of interpreting arthropod sampling with pitfall traps. Zoologischer Anzeig- er 202:177-184. Aitchison, C.W. 1978. Spiders active under snow in southern Canada. Symposia of the Zoological Society of London 42:139-148. Aitchison, C.W. 1984. The phenology of winter-ac- tive spiders. Journal of Arachnology 12:249- 271. Almquist, S. 1969. Seasonal growth of some dune- living spiders. Oikos 20:392-408. Berry, J.W. 1971. Seasonal distribution of common spiders in the North Carolina Piedmont. Ameri- can Midland Naturalist 85:526-531. Breymeyer, A. 1966. 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The spider population of a stand of oak in Wytham wood, Berks., England. The Canadian Entomologist 92:110-124. Uetz, G.W. 1977. Coexistence in a guild of wan- dering spiders. Journal of Animal Ecology 46: 531-541. Uetz, G.W. and J.D. Unzicker. 1976. Pitfall trapping in ecological studies of wandering spiders. Jour- nal of Arachnology 3:101-111. Williams, G. 1962. Seasonal and diurnal activity of harvestmen and spiders in contrasted habitats. Journal of Animal Ecology 31:21-42. Work, T.T., C.M. Buddie, L.M. Korinus, & J.R. Spence. 2002. Pitfall trap size and capture of three taxa of litter-dwelling arthropods: implica- tions for biodiversity studies. Environmental En- tomology 31:438-448. Manuscript received 19 September 2002, revised 29 May 2003, 2004. The Journal of Arachnology 32:231-269 A REVISION OF THE SPIDER GENUS CALILEPTONETA PLATNICK (ARANEAE, LEPTONETIDAE), WITH NOTES ON MORPHOLOGY, NATURAL HISTORY AND BIOGEOGRAPHY Joel M. Ledford: California Academy of Sciences, Department of Entomology, Golden Gate Park, San Francisco, California 94118, USA, and San Francisco State University, Department of Biology, 1600 Holloway Avenue, San Francisco, California 94132 USA. E-mail: jledford@comcast.net ABSTRACT, The spider genus Caiileptoneta Platnick is revised and all species are described, diagnosed and keyed. A neotype for Caiileptoneta californica (Banks) is designated and Caiileptoneta sylva (Cham= berlin & Ivie) is removed from synonymy with Caiileptoneta californica (Banks). The female of Calilep- toneta noyoana Gertsch and the male of Caiileptoneta sylva (Chamberlin & Ivie) are described for the first time. Three new species are described: Caiileptoneta briggsi, Caiileptoneta cokendolpheri, and Cai- ileptoneta ubicki. The morphology of Caiileptoneta is discussed, and interpretive illustrations of male and female genitalia are provided. The natural history of Caiileptoneta is presented with an account of the mating behavior for C. ubicki. The distribution of Caiileptoneta species is discussed, and areas of ende- mism and potentially new Caiileptoneta species are noted. Keywords; Taxonomy, new species, USA, Leptonetidae The first North American leptoeetid, Lep- toneta californica Banks, was described in 1908 and it was not until the time of Gertsch (1974) that the first effort to comprehensively treat the fauna was performed. Although de- tailed studies on the European fauna (Brignoli 1972, 1974, 1975, 1976, 1977, 1979a, 1979b, 1979c; Page 1913; Machado 1941, 1945) re- vealed great genitalic diversity, Gertsch relied largely on somatic characters, resulting in an inadequate description of the fauna and a se- ries of controversial taxonomic decisions (Brignoli 1977, 1979c; Platnick 1986). Plat- nick (1986) reassessed the higher-level tax- onomy of the North American leptoeetids by using a new suite of characters involving mid- dorsal integumentary glands and discovered support for the grouping of the North Amer- ican fauna into four genera, confirming the opinions of Brignoli (1977, 1979c). Little attention has since been paid to these spiders despite the problems found in Gertsch’s (1974) monograph. Most taxa suffer from incomplete species descriptions and doubtful species limitations. Additionally, over 50'% of the specimens studied by Gertsch (1974) are juveniles that cannot be diagnosed. This situation is especially troubling in west- ern North America, where the bulk of lepto- netid diversity is found in environmentally sensitive areas, and where a complete knowl- edge of these taxa may contribute to conser- vation efforts. The major systematic problem of the North American leptonetids is an incomplete under- standing of their genitalic morphology. A sur- vey of the European literature reveals a vari- ety of genitalic characters, especially on the male palpal bulb. None of the literature treat- ing the North American fauna sufficiently il- lustrates the complexity of the male genitalia, and the females are ignored almost entirely. Not only does this lack of information miss a suite of informative characters, it may also un- derestimate the true diversity of the fauna. This study reexamines the genus Caiilepto- neta Platnick 1986 and builds on the findings of Gertsch (1974) by using detailed genitalic examinations, natural history observations and additional specimens collected since his mono- graph. Three new species are described, in- cluding the first troglobitic Caiileptoneta. Nat- ural history observations are included, with the first account of the mating behavior of lepto- eetids and a description of their web architec- ture. The biogeography of Caiileptoneta is also discussed, noting areas of endemism and lo- calities that may yield additional species. It is 231 232 THE JOURNAL OF ARACHNOLOGY Table 1. — List of anatomical abbreviations used in the text and figures. AC apical constriction AEG anterior eye group AER anterior eye row ALE anterior lateral eyes ALS anterior lateral spinnerets AME anterior median eyes AH apical hook AL accessory lobe E embolus OA ocular area OAL ocular area length OAW ocular area width PBP proximal bulb process PEG posterior eye group PF proapical flange PL prolateral lobe PLE posterior lateral eyes PLS posterior lateral spinnerets PMS posterior median spinnerets PS paraembolar setae RS retroapical seta S spermathecae the aim of this study to provide a guideline for the revision of the other North American lep- tonetids, and contribute to an eventual under- standing of their phylogenetic history. METHODS Each species is thoroughly described, di- agnosed, and keyed. Species descriptions refer to a single adult individual for each sex, which is identified as a type or by the locality at which it was collected. Descriptions of fe- males and all previously unknown sexes were written using specimens collected in associa- tion with diagnosable individuals at or near the type locality. In cases of sympatry [C. cal- ifornica (Gertsch 1974) and C helferi Gertsch 1974)], descriptions were based upon associ- ated individuals collected as close to the type locality as possible. Anatomical abbreviations used in the text are listed in Table 1 . All mea- surements are in mm and quantify the size of a structure at its widest or longest point. A section reporting the variation in the most conspicuous and variable features follows each description and represents two to nine individuals (n), encompassing the full range in overall size. All illustrations are by Virginia Kirsch (VK) or myself (JL) and are attributed in the figure captions. Prior to examination with a Hitachi S-520 or Leo 1450 VP Scanning Electron Micro- scope, all structures were briefly cleaned in an ultrasonicator and critical point dried. Spin- neret preparations followed the methods of Coddington (1989), consisting of a brief cleaning in an ultrasonicator and a gentle squeeze of the abdomen using forceps in order to extend and separate the spinnerets. Large structures were examined using Olympus SZHIO and Leica SMZIO microscopes. Vulvae were carefully excised and placed in a trypsin solution for 2-8 hours under a heat lamp to digest extraneous tissue. If char- acters remained unclear, the vulva was stained with Chlorazol Black and reexamined. Pho- tographs were taken using an Olympus PM- lOAK and a Nikon Coolpix 990 digital cam- era attached to a Nikon SL3D® microscope. All genitalia and small structures were placed in Hoyer’s solution or glycerin and examined in temporary mounts following the procedure described by Coddington (1983), under a Ni- kon SL3D® microscope. Palpal structures are provided names based on the nomenclature used by Gertsch (1974) and Platnick (1986), when the structure re- ferred to was unambiguously identified. Struc- tures not previously discussed in the literature treating the North American fauna were given names based on their relative location on the palpal bulb. I did not use names provided in literature treating the European or Asian fau- nas. Such a system would imply homology of structures where none is intended. It is not the aim of this study to establish a precedent for the nomenclature of leptonetid genitalia, ex- cept in that structures should be clearly la- beled and illustrated. The specimens on which this study was based were kindly provided by Norman Plat- nick, American Museum of Natural History (AMNH), Laura Leipensburger, Museum of Comparative Zoology (MCZ), Fred Coyle, Western Carolina University (WCU), James Cokendolpher, personal collection (JC), and DaiTell Ubick, personal collection (DU). Ad- ditional material is from the collection of the California Academy of Sciences (CASC). SOMATIC MORPHOLOGY Cephalothorax. — The carapace (Figs. 8, 1 0) is domed and gently sloping posteriorly in lateral profile. The color is pale yellow with LEDFORD— REVISION OF CALILEPTONETA 233 Figure 1. — Web of Catileptoneta ubicki new species from Arroyo Seco Canyon. Scale bar = 2.0 mm. purple^browe dusky markings surrounding the eyes and carapace margins. The fovea is a lon- gitudinal purple-brown dusky band (Figs. 2, 4, 6). The eyes are relatively large (except in C briggsi new species), and raised above the carapace in lateral profile (Figs. 8, 10). Seta- tion is minimal, although a series of thin, elongate setae occur posteriad of the PME, and a distinct pair of crossing setae are at the apex of the clypeus (Figs. 2, 4, 6). The ster- num (Fig, 9) is slightly domed in lateral pro- file (Fig. 10) and the palpal coxae bear a ser- rula extending the length of the anterior edge (Fig. 11). The chelicerae (Figs. 15-23) have a promargin with a single row of large teeth on a fine ridge, and a retromargiri with 1-7 den- ticles. As with other leptonetids, the promar- gin bears a relatively large proximal tooth, al- though C. noyoana (Gertsch 1974) and C. wapiti (Gertsch 1974) males have a diagnostic enlarged tooth distally (Figs. 18, 19). Legs* — The legs are long and thin with a regular series of slender spines on the tibiae and metatarsi and a single apical spine on the patellae (Figs. 3, 5, 7). Sexual dimorphism of the legs is minimal, v/ith the leg length of males only slightly longer than that in females. A point of weakness occurs at the patella-tibia joint, resembling linyphiid, hersiliid and filis- tatid spiders, and the legs are readily dropped distad of this joint by living specimens (pers. obs.). A small preening comb consisting of 6 paired setae occurs ventroapically on the meta- tarsus (Figs. 12, 13). The comb is often tucked into the tibia-metatarsus joint and may require the removal of one of the segments to be clear- ly viewed. Similar combs have also been re- corded in Appaleptoneta and Neoleptoneta (Cokendolpher pers. comm.) and may be a syn- apomorphy uniting leptonetids as a whole. The integumentary glands discussed by Platnick (1986), occur middorsally on the femora, pa- tellae and tibiae of all legs, including the palpi. The sinuous patellar gland plates with large pores (Fig. 14) (Platnick, 1986 figs. 55-60) may indeed serve to unite Calileptoneta, al- though several other gland types were found on Calileptoneta species. No less than four dif- ferent types of glands were found on the legs of Calileptoneta sylva (Chamberlin & Ivie 1942), including gland types suggested as syn- apomorphies for other taxa by Platnick (1986) 234 THE JOURNAL OF ARACHNOLOGY Figure 2. — Calileptoneta ubicki new species, male, dorsal from Arroyo Seco Canyon. Scale bar = 0.5 mm. Illustration by VK. (Ledford & Ubick, pers. obs.). The tarsal claws are simple, with few teeth. Abdomen. — The abdomen (Figs. 2-7) is oval in shape, with several long, slender setae dorsally. The color is pale with a series of dusky chevron markings. These markings were used by Gertsch (1974) to diagnose spe- cies (especially juveniles) but I found them highly variable within species. Spinning organs. — The spinnerets (Figs. 24-31) of Calileptoneta are typical of other leptonetids and do not substantially differ from the findings of Platnick et. al. (1991). The ALS (Figs. 25, 30) are circular in apex outline, with 6-10 pustulose tartipores. The posterior spinnerets (Figs. 26, 27, 31) are highly modified, forming elongate, narrow tet- rahedrons bearing 1-2 rows of spigots with short, circular bases and narrow shafts. The PLS of females bear 1-2 elongate spigots with thick shafts (Fig. 31) that presumably serve the cylindrical glands (Platnick et. ah 1991). No such spigots were found in males, nor were any nubbins. A series of four epiandrous spigots (Fig, 28) occur at the apical edge of the epigastric furrow in males. GENITALIA Male. — The male genitalia (Figs. 32-94) of Calileptoneta are intricate structures that re- quire careful observation under compound mi- croscopy in order to be viewed properly. The palpal segments vary greatly in length, with some species (C helferi, C. noyoana) having elongate femora and patellae (Figs. 5, 7) sev- eral times the carapace width. The palpal tarsus bears an apical constriction (AC, Figs. 32, 35, 38) with a single seta retroapically (retroapical seta, RS, Figs. 33, 37, 39). Small groups of modified setae occur on the retrolateral surfac- es of the tibiae and tarsi (Figs. 37, 40, 68, 71, 86, 92). Spination is variable with C briggsi, C. californica, C. helferi and C sylva bearing numerous spines on the pro-and retrolateral surfaces of the palpal femora (Fig. 7), and C cokendolpheri new species, C. noyoana, C. oasa (Gertsch 1974), and C ubicki new spe- cies, lacking palpal spines entirely (Figs. 3, 5). The bulb is suboval in shape and lightly scler- otized. As reported by Brignoli (1979c), the bulb is expandable, and may rotate up to 90° during mating. An unusual prolongation of the palpal bulb (proximal bulb process, PBP, Figs. 38-40, 41-43, 47-49, 59-61, 77-79) extends proximally on C. briggsi, C. californica (Gertsch, 1974), C. helferi and C sylva, and may function as an anchoring point during mating. The prolateral lobe (PL, Figs. 32, 35, 38), first proposed as a synapomorphy for Cab ileptoneta by Platnick (1986), is a dorsal ex- LEDFORD— REVISION OF CALILEPTONETA 235 tension of the bulb (not the tarsus) that may serve as a point of rotation during bulb expan- sion. An accessory lobe (AL, Figs. 46, 82) oc- curs distad of the PL, and gradually tapers into a small hook at its apex (apical hook, AH, Figs. 35, 38, 46, 52, 56, 82, 87, 94). The ventroapical surface of the bulb is divided by a narrow ridge (Fig. 45) that is produced into a translucent, spoon-shaped, proapical flange with serrate edges (proapical flange, PF, Figs. 33, 36, 39). The prolateral surface of the ridge is deeply striate (Figs. 45, 46, 51, 52, 63, 64, 76, 82), composed of numerous laterally fused setae. The embolus (E, Figs. 33, 36, 39) is situated ventroapically and tapers into a point (Figs. 75, 76) or a broad fork (Fig. 46). An adjacent group of twisted circular or fan-like setae (par- aembolar setae, PS, Figs. 32, 34, 35, 38, 39) occur prolaterad of the embolus and may be the remnants of the laterally fused setae on the prolateral surface of the bulb. The tarsal organs (Figs. 95-97) of Calileptoneta are cylindrical structures situated at the apex of the palpal tar- sus, and vary in the degree by which they ex- tend from the tarsus. Female. — The female genitalia (Figs. 98- 118) of Calileptoneta are relatively uniform, and of little diagnostic value. Although many diagnostic characters were discussed by Gertsch (1974), most of his illustrations were drawn incorrectly and are misleading. The lack of diagnostic information in females was especially problematic during the course of this study, and all species except for C. cok- endolpheri, C. oasa and C. ubicki remain un- diagnosable in the absence of associated males. The atrium is triangular or trapezoidal with a dense network of striae (Figs. 102, 104, 106, 109-118) ventrally and a lateral pair of tightly twisted spermathecae (S, Figs. 98, 101) that connect dorsally via a sharp bend (Figs. 101, 107, 108). This bend is often difficult to observe and requires careful preparation to be viewed clearly (see Methods section for a dis- cussion of techniques that aid in the exami- nation of leptonetid genitalia). Two species, C. oasa and C. ubicki, have an atrium with a dis- tinctive apical bifurcation (AB, Figs. 98, 102, 104) although this is less pronounced in C. oasa. 236 THE JOURNAL OF ARACHNOLOGY Figure 4. — Calileptoneta noyoana (Gertsch), male, dorsal from Fort Bragg. Scale bar = 0.5 mm. Illustration by VK. NATURAL HISTORY General behavior and web architec- ture*— Calileptoneta are small spiders, gen- erally restricted to cool, moist microenviron- ments such as caves, leaf-litter and deeply imbedded rock outcrops. They are rarely col- lected due to their cryptic nature and temporal occurrence, especially in areas that are sea- sonally arid. Individuals appear to congregate in ideal habitats and, at the correct time of year, may be collected in large series. The web (Fig. 1) of Calileptoneta is a rectangular, fine- ly woven sheet 3-4cm in diameter, presum- ably constructed with the highly modified PLS. Individuals hang beneath the sheet and readily drop from the web into a defensive posture whereby the legs are drawn close to the body when disturbed (pers. obs.). Mating behavior* — A captive group of C ubicki were kept in a small plastic container containing sphagnum moss and fed a diet of freshly killed Drosophila for several months. Up to 7 individuals occupied the same con- tainer and overlapping sheet webs were con- structed with individuals often resting in close proximity. An adult pair of C ubicki was re- moved from the group and several instances of mating were recorded. Courtship displays consisted of a slight plucking of the female's web by the male using his palpi. Both indi- viduals remained inverted beneath the sheet as the male slowly approached. Upon contact, a single palpal bulb rotated 90° before being in- serted into the atrium and then exchanged for the other bulb. Alternate bulb insertion lasted approximately 5-10 seconds and mating was completed after 30 minutes. The male situated himself in the female’s web for several hours following mating. No sperm web was record- ed for the male and the female did not con- struct any eggsacs. BIOGEOGRAPHY The distribution of Calileptoneta (Figs. 119-122) is largely vicariant with most spe- cies restricted to a particular locality or habitat type. This is not surprising given their general biology and the similar patterns of endemicity exhibited by other leptonetid taxa (Brignoli 1974, 1977; Gertsch 1971). Sympatric popu- lations are known, however, and a strict hy- pothesis of vicariance may be premature given the poor sampling in central Oregon, eastern California and southern California (Fig. 122). The restriction of Calileptoneta species to specific habitat types is of special interest as environmental concerns are paramount in Cal- ifornia and Oregon. Of particular importance LEDFORD— REVISION OF CALILEPTONETA 237 Figure 5. — Calileptoneta noyoana (Gertsch), male, lateral from Fort Bragg. Scale bar =1,0 mm. Illus- tration by VK. are the distributions of C noyoana and C cokendolpheri, which are endemics of eco- nomically important forest types-redwood for- est and Douglas Fir forest, respectively. Al- though not particularly charismatic, nor common enough for use as indicators, these species contribute to the knowledge of biodi- versity in these forests and may assist conser- vation efforts. Troglobitic species regularly receive special conservation status due to their extremely lim- ited distributions and sensitivity to distur- bance. It is surprising that only a single trog- lobitic Calileptoneta is known, despite California’s rich cave fauna and in contrast to the other major karst regions in the Nearctic, Texas and Appalachia, which have numerous cavernicolus leptoeetids. In California, this niche appears to be occupied by the many cavernicolus telemids, which do not occur in the eastern United States (D. Ubick pers. comm.). Calileptoneta briggsi is restricted to a poorly known cave system that receives lit- tle attention from cavers and is probably not at risk. Additional troglobitic Calileptoneta al- most certainly occur, and indeed two female specimens have been collected in caves from Caleveras and Tulare counties in eastern Cal- ifornia (Fig. 122). Given the large disjunction between these specimens and other known populations of Calileptoneta, they almost cer- tainly represent new species. Undiagnosable females and juveniles (Fig. 122) provide additional interesting distribu- tional data for Calileptoneta. Not only do these specimens contribute to a broader un- derstanding of the distribution of Calilepto- neta, they reveal distributional gaps that may yet yield additional species. The gap in south- ern California is the most striking, with only two species, C. ubicki and C. oasa, known from the entire southern half of the state. The seasonally arid climates of these areas makes the collection of specimens unpredictable, but given the habitat diversity and faueistic rich- ness of these areas, the possibility of this be- ing a natural distributional gap is unlikely. The large gap in Oregon from Josephine County to Lane County is also probably arti- ficial considering the rich oak woodland and riparian areas that occur betv^eee these two counties. 238 THE JOURNAL OF ARACHNOLOGY Figure 6. — Caiileptoneta helferi (Gertsch), male, dorsal from Mt. Diablo. Scale bar = 0.5 mm. Illus- tration by VK. DISCUSSION Perhaps the largest remaining gap in the knowledge of the North American leptonetids is an understanding of their phylogentic affin- ities. Not only would a phylogeny for these spiders reveal interesting evolutionary pat- terns within North America, it would also contribute to an understanding of leptonetid relationships as a whole. Such a study would require a comprehensive review of all North American leptonetid genera, and inclusion of potential European and Asian outgroup taxa, which is beyond the scope of this study. The relationships within Caiileptoneta also need further investigation and future work will be directed at this goal. Additional fieldwork is prerequisite to this understanding, especially in the distributional gaps of central Oregon, southern California and eastern California. Additional specimens will also facilitate the diagnosis of females, particularly between C. briggsi, C. californica, C. helferi and C. noy- oana. The recollection of C. wapiti also must take priority, as will the discovery of males at cave sites in Calaveras and Tulare counties. TAXONOMY FAMILY LEPTONETID AE Genus Caiileptoneta Platnick 1986 Leptoneta (in part); Banks 1910: 6; Brignoli 1977: 215-217; Comstock 1913: 307; Page 1913: 566; Moles 1921: 40; Gertsch 1974: 191-192. Caiileptoneta Platnick 1986: 15; Platnick 2002. Type species. — Leptoneta californica Banks 1904: 333, by original designation. Diagnosis. — Distinguished from other North American leptonetid genera (Archolep- toneta Gertsch 1974, Neoleptoneta Brignoli 1972, Appaleptoneta Platnick 1986) by having a prolateral lobe on the male palpal tarsus (Figs. 32, 35, 38) and by the sinuous patellar gland plates bearing large pores (Fig. 14, Plat- nick 1986, figs, 55-60). Description. — Total length 1.20-3.04. Car- apace oval in dorsal view (Fig. 8), length 1.20-1.32 X width, height 0.26-0.54 X width; smooth; thoracic fovea a thin dusky band (Figs. 2, 4, 6); carapace posterior margin straight to sinuous (Figs. 2, 4, 6, 8); carapace with 2 setae posteriad of AER, 3 setae pos- teriad of PME, extending linearly to fovea; clypeus with 2 setae crossing distally (Figs. 2, 4, 6); six eyes, PME absent; AEG with dark markings surrounding eyes, u-shaped at pos- terior margin (Figs. 2, 4, 6, 8); PEG with dark circular markings surrounding eyes (Figs. 2, 4, 6); OA longer than wide, OAL 0.60-1.40 X OAW, PME 0.40-0.70 X PLE interdisL ances; clypeal height 0.70-1.02 X PME di- ameter, chelicerae unmodified, fang furrow a narrow ridge with 3-9 large teeth, retromargin with 1-5 denticles (Figs. 15-23). Sternum smooth, oval, broadly rounded posteriorly, margin entire, length L0-L2 X width. Ab- domen (Figs. 2, 4, 6) oval, pale, with dusky LEDFORD— REVISION OF CALILEPTONETA 239 chevron markings, and covered with fine, elongate setae. Spinning organs (Figs. 24-31) with the ALS not modified and bearing sev- eral pustulose tartipores; PMS and PLS highly modified, tetrahedral, forming a narrow ridge on surface; PMS with 2 longitudinal rows, and PLS with a single longitudinal row of spigots on short, circular bases; prolateral edge of PLS in females with 2 isolated spigots on elongate, thick bases (Fig. 31); males with 4 epiandrous spigots (Fig. 28) at the apex of the epigastric furrow. Leg formula I, IV, III, II; elongate and thin; femur I of males 2.09-3.4 X carapace width, females 1.32-2.71 X car- apace width; middorsal integumentary glands sinuous, with large pores (Fig. 14); preening comb consisting of 6 paired setae at the apices of the metatarsi on legs I~IV (Figs. 12, 13); autospasy occurs at patella- tibia joint. Leg spi- nation I-IV: patella dl, tibia dl-1, pi, rl. Male palpal femur length 0.67-2.78 X cara- pace width; palpal tarsus with a retroapical seta (Figs. 33, 37, 39). Bulb expandable, light- ly sclerotized, suboval, longer than wide, length 0.40-2.20 X tibia length; with a fine ridge ventroapically, and a transparent proap- ical flange with serrate edges (Figs. 33, 36, 39); prolateral surface bearing a large lobe (Figs. 32, 35, 38), with a small lobe apicad (Figs. 46, 82) that is produced distally into a hook (Figs. 35, 38, 46, 52, 56, 64, 82, 87, 94); embolus ventrally situated (Figs. 33, 36, 39), with a prolateral group of twisted circular or fan-like paraembolar setae (Figs. 32, 34, 35, 38, 39); tarsal organ (Figs. 95-97) cylindrical. Female genitalia (Figs. 98-118) with a trian- gular or trapezoidal atrium, length 0.72-1.12 X width, densely striate on ventral surface; spermathecae situated laterally, elongate, twisted, attaching to atrium dorsally by a sharp lateral bend (Figs. 101, 107, 108). Distribution..— Southern California (Riv- erside County) to Central Oregon (Lane County) (Figs. 119-122). KEY TO SPECIES OF THE GENUS CALILEPTONETA The key has been written to maximize accuracy and efficiency. Multiple features are pre- sented in each couplet in order to assist the identification of variable and poorly preserved specimens. Most taxa require high magnification under compound microscopy to insure pos- itive identification. Males 1. Proximal bulb process present (Figs. 38-40), retroapical seta straight (Figs. 33, 39), pro- and retrolateral surfaces of palpal femur heavily spined (Fig. 7) ..................... 2 Proximal bulb process absent (Figs. 32-37), retroapical seta straight to curved (Fig. 37), palpal femur weakly spined or lacking spines entirely (Figs. 3, 5).. ......... ......... 5 2. Proximal bulb process elongate, length 1.0-2.04 X bulb width (Figs. 47-49, 59, 61), ac- cessory lobe reduced (Figs. 52, 64) ............................................ 3 Proximal bulb process shorter, length 0.55-1.0 X bulb width (Figs. 41-43, 77-79), accessory lobe normal (Figs. 46, 82) ................................................... 4 3. Palpal tibia elongate, length 1.02-1.19 X carapace width, proximal bulb process extending length of tibia, bulb length 0.81-1.04 X tibia length (Figs. 6-7, 59-64), body darkly pigmented ................................................... C helferi (Gertsch) Palpal tibia shorter, length 0.76-0.90 X carapace width, proximal bulb process extending entire length of tibia, bulb length 1.09-1.40 X tibia length (Figs. 47-52), body lightly pigmented ................................................. C. californica (Banks) 4. Body pigmentation entirely lacking, eyes reduced and flattened, femur I elongate, length 2.88-3.40 X carapace width; proximal bulb process longer, length 0.85-1.0 X bulb width (Figs. 41-43), embolus apically forked (Fig. 46) ................. C. briggsi new species Body darkly pigmented, eyes normal, diameter PME 0.50-0.64 X PLE interdistances; prox- imal bulb process shorter, with slight retrolateral bend (Fig. 78), length 0.55-0.86 X bulb width (Fig. 79) ....................................... C. syiva (Chamberlin & Ivie) 5. Palpal tarsus with a retrolateral pair of twisted setae (Figs. 37, 65-70, 92), chelicerae with an enlarged distal tooth (Figs. 18, 19), retrodistal chelicerai process absent 6 240 THE JOURNAL OF ARACHNOLOGY Palpal tarsus lacking twisted setae, chelicerae without an enlarged distal cheliceral tooth (Figs, 15-17, 20-23), retrodistal cheliceral process present (Fig. 16) or absent .......... 7 6. Palpal segments elongate, femur length 2.24—2.78 X carapace width (Figs. 5, 65-70) .... ........................................................... C. noyoana (Gertsch) Palpal segments normal, femur length 0.63-1.0 X carapace width (Fig. 89-94) ......... ............................................................ C wapiti (Gertsch) 7. Bulb bearing two proapical flanges; proximal flange setose, distal flange tightly curled (Fig. 54); embolus hook-shaped (Fig. 56) ...................... C. cokendolpheri new species Bulb with single proapical flange, lacking setae, apex loosely to tightly curled (Figs. 75, 76, 88), embolus tapering to fine point (Figs. 75, 76) ................................. 8 8. Retrodistal cheliceral apophysis present (Fig. 16), with a whip-shaped seta retroapically on the palpal tibia (Fig. 86), proapical flange sinuate (Fig. 88) .......... C. ubicki new species Retrodistal cheliceral apophysis absent (Fig. 15), with a hook-shaped seta retroapically on the palpal tibia (Fig. 71), proapical flange straight (Fig. 75) ............. C. oasa (Gertsch) Females 1. Atrium with an apical bifurcation (Figs. 98, 102, 104) .......................... ........................................... C oasa (Gertsch), C ubicki new species Atrium entire (Figs. 99, 100, 106, 109-1 18) ..................................... 2 2. Atrium (Figs. 99, 100, 106, 109-116) distinctly triangular, tapering to a point apically, with normal base .............................................................. . . , C. briggsi new species, C. californica (Banks), C. helferi (Gertsch), C. noyoana (Gertsch), C. wapiti (Gertsch) Atrium (Fig. 117-118) trapezoidal, apically subquadrate, with broad base, atrium length 0.72-0.81 X width ................................... C, cokendolpheri new species Calileptoneta briggsi new species Figs. 22, 41-46, 111, 112, 121 Type material. — Male holotype from In- dian Valley Creek Cave, Trinity County, Cal- ifornia, USA, 40°37'N, 123°27'W, 27 October 1990, D. Ubick, W. Rauscher (CASC). Para- types: USA: California: 26, 5 $, same data as holotype (CASC; 1 6, 1 $, DU). Other material examined. — USA: Cali- fornia: Trinity County: Indian Valley Creek Caves, 4 air miles SSE. Hyampom, 40°37'N, 123°27'W, -1800 ft elevation, 5 July 1980, X Briggs, W. Ma, W. Rauscher (29,1 juve- nile, AMNH), 27 October 1990, D. Ubick, W. Rauscher (4 juveniles, CASC), 31 August 1996, D. Ubick (2 juveniles, DU); Lower But- ter Creek Cave, 40°37'N, 123°27'W, 5 July 1980, T. Briggs, W. Ma, W. Rauscher (2 9,4 juveniles, AMNH). Etymology. — This species is named in honor of Dr. Tom Briggs, discoverer of this and many other troglobitic arachnids through- out California. Diagnosis. — Distinguished from all other Calileptoneta by lacking pigmentation and having the eyes greatly reduced and flattened. Calileptoneta briggsi may be further separated from other Calileptoneta males, except C cal- ifornica, C. helferi and C. sylva, by having a proximal bulb process (Figs. 38-43), and a straight retroapical seta (Figs. 39, 48); from C californica and C helferi by having the proximal bulb process (Figs. 77-79) short, process length 0.85-1.08 X bulb width, and having the prolateral apical lobe large (Figs. 46, 82); from C sylva by having an elongate femur I, length 2.88-3.40 X carapace width, and an apically forked embolus (Fig. 46). Male (holotype), — Total length 2.64. Spec- imen pale, entirely lacking pigmentation. Car- apace 1.19 long, 0.93 wide, height at fovea 0.33 X carapace width; clypeus 0.17 high, chelicerae 0.66 long, fang furrow with 9 teeth on a narrow ridge and 5 denticles on retro- margin (Fig. 22). Ocular area 0.23 long, 0.17 wide; diameter PME 0.50 X PLE interdist- ances. Sternum 0.65 long, 0.17 wide; labium 0,08 long, 0.16 wide; palpal coxae 0.54 long, 0.20 wide. Spination: palpus: femur pl-1-2-1-2-2-2-2- 3-4-1, r3-2-l-l-2-3-2; patella dl; tibia r3; tar- sus rl (apical). Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [To- LEDFORD— REVISION OF CALILEPTONETA 241 Figure 7. — Calileptoneta helferi (Gertsch), male, lateral from Mt. Diablo. Scale bar =1.0 mm. Illus- tration by VK. tal]): I: 3.17 + 0.39 + 2.80 + 2.24 + 1.34 = [9.95]; II: 1.90 + 0.34 + 2.04 4- 1.73 + 1.10 - [7.12]; III: 1.70 + 0.27 + 1.59 + 1.46 + 0.90 = [5.90]; IV: 2.10 + 0.32 + 2.15 + 1.93 + 1.15 = [7.65]; pedipalpus: 1.08 + 0.44 + 0.66 + 0.41 = [2.60]. Femur I 3.40 X cara- pace width, palpal femur 1.16 X carapace width. Palpal bulb (Figs. 41-46) 0.79 long, 0.27 wide; palpal tibia with a retroapical group of stiff setae; proximal bulb process (Figs. 41- 43) short, reaching to base of tibia, bulb length 0.98 X length tibia; embolus broadly forked at apex (Fig. 46); paraembolar setae circular, distally broad, reaching slightly be- yond base of fork on embolus (Fig. 46); ac- cessory lobe large (Fig. 46). Abdomen pale, without pattern, 1.45 long, 1.22 wide. Variation {n = 2). — Total length 2.05- 2.64; carapace length 1.24-1.31 X carapace width; OAL 1.35-1.77 X OAW, diameter PME 0.40-0.53 X PLE interdistances; length femur I 2.10-3.40 X carapace width, palpal femur 0.91-1.20 X carapace width; palpal bulb length 1.15-1.36 X palpal tibia length; proximal bulb process length 0.85-1.08 X bulb width. Female (paratype). — Total length 2.90. Coloration and markings same as male. Carapace 1.2 long, 0.93 wide, height at fo- vea 0.34 X carapace width; clypeus 0.17 high, chelicerae 0.65 long, fang furrow with 8 teeth on a narrow ridge and 4 denticles on retro- margin (Fig. 22). Ocular area 0.11 long, 0.18 wide; diameter PME 0.50 X PLE interdist- ances. Sternum 0.77 long, 0.70 wide; labium 0.14 long, 0.20 wide; palpal coxae 0.49 long, 0.23 wide. Spination: palpus: patella dl (apical), tarsus p3-l-l, rl-4, vl. Leg measurements (Eemur + Patella + Tibia + Metatarsus + Tarsus = [To- tal]): I: 2.32 + 0.34 + 2.93 + 1.98 + 1.27 - [8.84]; II: 1.93 + 0.34 + 1.98 + 1.63 + 1.02 = [6.9]; III: 1.63 + 0.29 F 1.49 + 1.46 + 0.90 = [5.77]; IV: 2.15 + 0.29 + 2.10 + 1.88 242 THE JOURNAL OF ARACHNOLOGY Figures 8-1 L — Calileptoneta sp., female from Mt. Diablo. 8. Carapace, dorsal. 9. Cephalothorax, ven- tral. 10. Cephalothorax, lateral. 11. Right palpal coxa showing serrula. Scale bars: A-C = 270 jxm, D = 20 fxm. + L12 = [7.54]; pedipalpus: 0.80 + 0.23 + 0.61 + 0.70 = [2.34]. Femur I 2.50 X cara- pace width, palpal femur 0.86 X carapace width. Abdomen pale, without pattern, 1.7 long, 1.35 wide. Atrium 0.18 long, 0.23 wide, sper- mathecae 0.18 long (Figs. 111-112). Variation (w ” 3). — Total length 2.63- 3.04; carapace length 1. 26-1. 32 X carapace width; OAL 0.60-1.45 X OAW, diameter PME 0.40-0.50 X PLE interdistances; length femur I 2.5-2.71 X carapace width, palpal fe- mur 0.82-0.89 X carapace width; atrium length 0.78-0.91 X width, spermathecae 0.78-0.80 X atrium width. Natural history. — These spiders were col- lected hanging from sheet webs among a root mass in the cave's dark zone. Given the lack of pigmentation, reduced eyes, and elongate legs of this species, it is considered a troglob- ite. Distribution. — Known only from Indian Valley Creek Caves which include Indian Val- ley Creek Cave and the adjacent Lower Butter Creek Cave in Trinity County (Eig, 121). These caves occur along a continuous band of LEDFORD— REVISION OF CALILEPTONETA 243 Figures 12-14. — Calileptoneta sp., female from Mt. Diablo. 12-13. Left metatarsus III, showing preen- ing comb. 14. Middorsal integumentary gland, left leg III. Scale bars: A = 150 p.m, B = 38 p-m, C = 7.5 pm. limestone and were presumably once con- nected (T. Briggs, D. Ubick pers. comm.). Calileptoneta californica (Banks 1904) Figs. 20, 38-40, 47-52, 115, 116 Leptoneta californica Banks 1904: 333; Banks 1910: 6; Brignoli 1977: 217; Comstock 1914: 307; Page 1913: 566; Moles 1921: 40; Gertsch 1935: 21; Gertsch 1974: 191-192. Calileptoneta californica (Banks): Platnick 1986: 15. Platnick 2002. Type material. — Female holotype, Mt. Diablo, Contra Costa County, California, 244 THE JOURNAL OF ARACHNOLOGY Figures 15-23. — Calileptoneta species, left chelicerae, female, male. 15. C. oasa (Gertsch) from An- dreas Canyon. 16. C. ubicki new species from Arroyo Seco Canyon, arrow to retrodistal cheliceral apoph- ysis. 17. C. cokendolpheri new species from H. J. Andrews. 18. C. wapiti (Gertsch) holotype, arrow to distal tooth. 19. C. noyoana (Gertsch), female from Fort Bragg, male holotype, arrow to distal tooth. 20. C. californica (Banks), female from Bell Station, male from Mt. Diablo. 21. C. helferi (Gertsch), female from Claremont Ave., male holotype. 22. C. briggsi new species from Indian Valley Creek Cave. 23. C. sylva (Chamberlin & Ivie), female holotype, male from Samwell Cave. Illustrations by JL. USA, 37°51'N, 121°55'W, June, Fuchs (CASC, lost in 1906 fire). Male neotype, Mt. Diablo State Park, BBQ/ Wildcat Group Camp, Contra Costa County, California, USA, 37°51'N, 121°55'W, 22 January 2000, J.M. Ledford, under stones (CASC); Other material examined. — USA: Cali- fornia: Contra Costa County: Mt. Diablo State Park, BBQ/ Wildcat Group Camp, 37°5UN, 121°55'W, 22 January 2000, J.M. Ledford, un- der stones (1 6, CASC); Napa County: 2 miles W Oakville, 38°26'N, 122°24'W, 31 Dec. 1953, V. Roth (1 d, 1 $, AMNH); Santa LEDFORD— REVISION OF CALILEPTONETA Clara County: 9.0 miles N. Bell Station, 37°02'N, 121°18'W, oak grove, under schist, 10 February 1991, D. Ubick (1 d, 2 $, DU). Designation of neotype. — The holotype of L. californica was lost in the California Acad- emy of Sciences during the 1906 earthquake and fire. Many additional specimens have since been collected at the type locality, Mt. Diablo. However, this locality represents an area of sympatry between C californica and C helferi (Fig. 120). Calileptoneta californica is by far the rarest of the two species with only two male specimens known from this locality. Gertsch (1974) assigned males with the palpal bulb process reaching the tibia (Figs. 47-52) to C. californica and I maintain the associa- tion. In order to define this species objectively and clarify its taxonomic status a male from Mt. Diablo conforming to both Banks' (1904) and Gertsch’s (1974) description of C calE fornica is designated as a neotype and depos- ited at the California Academy of Sciences (ICZN 2000, Article 28). Diagnosis.— Distinguished from other Cal- ileptoneta, except C. briggsi, C helferi and C sylva, by males having a proximal bulb pro- cess (Figs. 38-40, 47-49), and a straight re- troapical seta (Figs. 39, 48); from C. briggsi and C. sylva by having the proximal bulb pro- cess (Figs. 47-49) elongate, process length 1.0-2.04 X bulb width and having the acces- sory lobe reduced (Fig. 52); from C helferi by having a shorter palpal tibia, length 0.76- 0.90 X carapace width, whereby the proximal bulb process extends the entire length of the tibia, bulb length 1.09-1.40 X tibia length (Figs. 47-49). Male (eeotype).- — ^Total length 1.93. Cara- pace pale yellow-brown with fine dusky mot- tling surrounding margin, and laterally along caput margins; clypeus with dusky mottling distally; sternum dusky; coxae, trochanters, legs, and pedipalpi with dusky mottling, being especially conspicuous at the bases and apices of the segments. Carapace 0.80 long, 0.65 wide, height at fo- vea 0.30 X carapace width; clypeus 0.11 high, chelicerae 0.41 long, fang furrow with 7 teeth along a narrow ridge and 3 denticles on retro- margin (Fig. 20). Ocular area 0.20 long, 0.16 wide; diameter PME 0.60 X PLE interdist- ances. Sternum 0.52 long, 0.47 wide; labium 0.06 long, 0.12 wide; palpal coxae 0.34 long, 0.16 wide. 245 Spination: palpus: femur pi -2-1 -2-2-2, r2- 2-2-2-2-1 (apical), patella dl, tibia r3, tarsus rl. Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]): I: 1.64 + 0.26 + 1.81 + 1.51 + 0.97 = [6.19]; II: 1.23 + 0.26 + 1.21 + 1.05 + 0.75 - [4.50]; III: 1.01 + 0.24 + 0.91 + 0.91 + 0.63 = [3.70]; IV: 1.37 + 0.26 + 1.31 + 1.21 + 0.79 = [4.94]; pedipalpus: 0.86 + 0.71 + 0.99 + 0.55 = [3.11]. Femur I 2.52 X carapace width, palpal femur 1.32 X carapace width. Palpal bulb (Figs. 47-52) 0.69 long, 0.22 wide; palpal tibia with a retroapical group of stiff setae; proximal bulb process (Figs. 47- 49) elongate, reaching to base of tibia, bulb length 0.90 X length tibia; embolus narrowly forked at apex; paraembolar setae circular, reaching to base of fork on embolus (Figs. 51, 52); prolateral apical lobe reduced (Fig. 52). Abdomen dusky with pale chevron pattern, 1.13 long, 0.76 wide. Variation (#i — 4). — ^Total length 1.86- 1.93; carapace length 1.11-1.23 X carapace width; OAL 1 16-1.35 X OAW, diameter PME 0.55-0.64 X PLE interdistances; length femur I 2.24-2.52 X carapace width, palpal femur 1.25-1.42 X carapace width; palpal bulb length 1.09-1.40 X palpal tibia length; proximal bulb process length 1.0-1.54 X bulb width. Female (Bell Station).— Total length 2.31. Coloration and markings same as male. Carapace 0.91 long, 0.72 wide, height at fo- vea 0.26 X carapace width; clypeus 0.13 high, chelicerae 0.49 long, fang furrow with 7 teeth on a narrow ridge and 5 denticles on retro- margin (Fig. 20). Ocular area 0.20 long, 0.18 wide; diameter PME 0.50 X PLE interdist- ances. Sternum 0,54 long, 0.54 v/ide; labium 0.06 long, 0.16 wide; palpal coxae 0.37 long, 0.17 wide, Spination: palpus: patella dl, tibia rl-1, tar- sus p3-l (apical), rl (apical), vl (apical). Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus [Total]): I: 1.76 + 0.32 + 1.96 + 1.58 + 1.05 - [6.67]; II: 1.41 + 0.28 + 1.43 + 1.13 + 0.79 - [5.04]; III: 1.23 + 0.26 + 1.05 + 1.01 + 0.46 - [4.01]; IV: 1.61 + 0.26 + 1.58 + 1.37 + 0.87 = [5.69]; pedipalpus: 0.61 + 0.19 + 0.42 + 1.01 = [2.23]. Femur I 2.44 X carapace width, palpal femur 0,85 X carapace width. Abdomen 1.40 long, 1.12 wide. Atrium 246 THE JOURNAL OF ARACHNOLOGY LEDFORD— REVISION OF CALILEPTONETA 0.17 long, 0.22 wide, spermathecae 0,73 long (Fig. 113). Variation (n — 2). — Total length 2.05- 2.31; carapace length 1.26-1.27 X carapace width; OAL LI 1-1.18 X OAW, diameter PME 0.50-0.60 X PLE ieterdistaeces; length femur I 2.40-2.44 X carapace width, palpal femur 0.80-0.85 X carapace width; atrium length 0.77-0.83 X width, spermathecae 0.61-0.73 X atrium width (Fig. 114). Natural Mstory,— The Mt. Diablo speci- mens were found under moist stones in oak woodland sympatrically with C. helferi. This species appears to be restricted to drier habi- tats, unlike C helferi, which also occurs in redwood forest. Interestingly, a single speci- men sifted from redwood duff along the Smith River in northern California (Fig. 120) has a palp that conforms to C. caiifornica, however, considering its badly damaged condition (al- most nothing of the spider remains except the palp), and unknown collector, it is placed as C. californica incertae sedis. Distribution.— Central and northwestern California (Fig. 120). Calileptoneta cokemdolpheri new species Figs. 17, 53-58, 117-118, 121 Type materiaL— Male holotype from pit- fall traps in old growth Douglas Fir at the Uni- versity of Oregon’s H. J. Andrews Experi- mental Forest, Lane County, Oregon, USA, 44°10'N, 122n9'W, June-July 1987, no coL lector listed (AMNH). Paratypes: USA: Oregon: same data as holotype (6 d, 3 ?, AMNH). Etymology.— This species is named in honor of Mr. James Cokendolpher, who con- tributed additional leptonetid specimens to this study and shared insights into the often- times difficult morphology of these spiders. Diagnosis.- — Males are distinguished from all other Calileptoneta species by having 2 proapical flanges (Fig. 57), with the basal flange bearing numerous setae, and the apical flange being tightly curled. Calileptoneta cok- endolpheri may be further distinguished from other Calileptoneta, except C. oasa and C 247 ubicki, by lacking a proximal bulb process (Figs. 32-34) and the retroapical pair of twist= ed palpal tarsal setae; and from C oasa and C. ubicki by having a hook=shaped embolus (Fig. 56). Male (holotype). — Total length 2.33. Spec- imen pale. Carapace and all leg segments, in- cluding pedipalps, dark yellow-brown. Carapace 1.10 long, 0.91 wide, height at fo- vea 0.50 X carapace width; clypeus 0.19 high, chelicerae 0.93 long, fang furrow with 9 teeth along a narrow ridge and 4 denticles on retro- margin (Fig. 17). Ocular area 0.15 long, 0.19 wide; diameter PME 0.64 X PLE ieterdisL ances. Sternum 0.60 long, 0.65 wide; labium 0.11 long, 0.18 wide; palpal coxae 0.52 long, 0.22 wide. Spinatioe: palpus: patella dl, tibia rULl- L2 (apical), tarsus rl (apical). Leg measure- ments (Femur + Patella + Tibia L Metatarsus + Tarsus == [Total]): I: 1.88 + 0,32 + missing + missing + missing = [N/A]; II: 1.46 + 0.29 + 1.51 + 1.34 + 0.85 = [5.45]; III: 1.24 + 0.29 + 1.24 + 1.15 + 0.78 = [4.70]; IV: 1.63 + 0,29 + missing + missing + missing = [N/ A]; pedipalpus: 0.54 + 0.23 + 0.29 + 0.35 = [1.41]. Femur I 2.07 X carapace width, palpal femur 0.59 X carapace width. Palpal bulb (Figs. 53-58) 0.42 long, 0.23 wide; palpal tibia with a retroapical group of stiff setae; embolus hook- shaped (Fig. 56); paraembolar setae fae-like, with a single seta extending to base of hook on embolus (Fig. 56); ventral ridge bearing 2 proapical flanges (Fig. 57), with the basal flange bearing nu- merous setae, and the apical flange being tightly curled; prolateral apical lobe reduced (Fig. 58). Abdomen dark, without chevron pattern, 1.23 long, 1.07 wide. Variation (n = 2).— Total length 2.14- 2.33; carapace length 1.20-1.22 X carapace width; OAL 0.78-1.22 X OAW, diameter PME 0.64 X PLE ieterdistances; length femur I 2.07-2.15 X carapace width, palpal femur 0.59-0.76 X carapace width; palpal bulb length L30-L46 X palpal tibia length. Figures 24-28. — Calileptoneta helferi (Gertsch), male from Mt. Diablo, spinning organs. 24. Ventral. 25. Left ALS, arrow to tartipore. 26. Right PMS. 27. Right PLS. 28. Epiandrous spigots. Scale bars: A = 120 |xm, B = 13.6 fxm, C = 27 pm, D = 25 pm. 248 THE JOURNAL OF ARACHNOLOGY Figures 29-31. — Calileptoneta sp., female from Mt. Diablo, spinning organs. 29. Ventral. 30. Left ALS, arrow to tartipore. 31. Left PLS, PMS, anows to cylindrical gland spigots. Scale bars: A = 120 jjum, B = 13.6 (xm, C = 30 p.m. Female (paratype). — Total length 2.47. Coloration and markings same as male. Carapace 1.0 long, 1.0 wide, height at fovea 0.45 X carapace width; clypeus 0.19 high, chelicerae 0.58 long, fang furrow with 9 teeth along a narrow ridge and 5 denticles on retro- margin (Fig. 17). Ocular area 0.21 long, 0.19 wide; diameter PME 0.50 X PLE interdist- ances. Sternum 0.62 long, 0.65 wide; labium 0.09 long, 0.17 wide; palpal coxae 0.51 long, 0.21 wide. Spination: palpus: patella dl, tibia vl-2, tar- sus p 1 (apical), rl (apical), v 1 (apical). Leg measurements (Femur -f Patella + Tibia + LEDFORD— REVISION OF CALILEPTONETA 249 Figures 32-40. — Calileptoneta sp., male genitalia, left palpus. 32-34, C, oasa, Andreas Canyon. 35- 37 C. wapiti, Cameron Road, arrows on 37 to retroapical tibial setae. 38-40. C. californica Mt. Diablo, arrows on 40 to retroapical setae. AC = apical constriction, AH = apical hook, AL = apical lobe, E = embolus, PBP = proximal bulb process, PF = proapical flange, PL = prolateral lobe, PS == paraembolar setae, RS = retroapical tarsal seta, T = tibia, t = tarsus. Scale bars: 32-34 = 0.20 mm, 35-37 = 0.20 mm, 38-40 = 0.40 mm. Illustrations by JL. 250 THE JOURNAL OF ARACHNOLOGY Figures 41-46. — Calileptoneta briggsi new species, male from Indian Valley Creek Cave, right palpus. 41. retrolateral. 42. ventral. 43. prolateral. 44. retrolateral. 45. ventral. 46. prolateral. AC = apical con- striction, AH = apical hook, AL = apical lobe, E = embolus, PBP = proximal bulb process, PF = proapical flange, PL = prolateral lobe, PS = paraembolar setae, RS = retroapical setae. Scale bars: 41- 43= 250 |jLm, 44-46 = 100 ixm. Metatarsus + Tarsus = [Total]): I: 1.78 + 0.32 + missing + missing T missing = [N/A]; II: 1.46 + 0.27 + 1.46 + 1.15 + 0.80 = [5.14]; III: 1.27 + 0.24 + missing + missing + miss- ing = [N/A]; IV: 1.54 + 0.24 + 1.32 + 1.29 + missing = [N/A]; pedipalpus: 0.53 + 0.20 + 0.40 + 0.59 = [1.72]. Femur I 1.78 X car- apace width, palpal femur 0.53 X carapace width. Abdomen 1.47 long, 0.96 wide. Atrium 0.18 long, 0.25 wide, spermathecae 0.17 long (Fig. 117). Variation {n = 2). — Total length 2.44- 2.47; carapace length 1.0-1.27 X carapace width; OAL 1.0-1.10 X OAW, diameter PME 0.50 X PLE interdistances; length femur I 1.54-1.78 X carapace width, palpal femur 0.51-0.53 X carapace width; atrium length 0.72-0.81 X width, spermathecae 0.68-0.69 X atrium width (Fig. 118). Natural history, — The entire series of specimens representing this species were col- lected in old growth Douglas Fir {Pseudotsu- ga menziesii) using pitfall traps. Distribution. — Known only from the type locality (Fig. 121). Calileptoneta helferi (Gertsch 1974) Figs. 6, 7, 21, 24-28, 59-64, 115-116, 120 Leptoneta helferi Gertsch 1974: 192-194; Brignoli 1977: 217. Calileptoneta helferi (Gertsch): Platnick 1986: 15; Platnick 2002. Type material. — Male holotype, Carlotta, Humboldt County, California, USA, 40°32'N, 124°03'W, 15 September 1961, W. Ivie, W.J. Gertsch (AMNH, examined). Other material examined. — USA: Cali- fornia: Alameda County: Claremont Ave, 2.2 miles above Berkeley RB-2, 37°52'N, 122°16'W, 10 May 1963, PR. Craig, D. Dailey (1 $, 1 (3, CASC); Contra Costa County: Mt. LEDFORD— REVISION OF CALILEPTONETA 251 Figures 47-52. — Calileptoneta califomica (Banks), male from Mt. Diablo, right palpus. 47. retrolateral. 48. ventral. 49. prolateral. 50. ventroapical. 51. ventral. 52. prolateral. AC = apical constriction, AH = apical hook, AL = apical lobe, E = embolus, PBP = proximal bulb process, PF = proapical flange, PL = prolateral lobe, PS = paraembolar setae, RS = retroapical setae. Scale bars; 47 = 150 |xm, 48-49 = 231 |xm, 50 = 75 fxm, 51-52 = 86 jxm. Diablo State Park, BBQ/ Wildcat Group Camp, 37°5rN, 12r55'W, 22 January 2000, J.M. Ledford, under stones (4 d, 3 $, CASC), 6 February 2000, J.M. & K.E. Ledford, under stones (3 d, 2 $, CASC); Humboldt County: F.K. Lane State Park., nr. Phillipsville, 40°12'N, 123°47'W, 1 Oct 1959, V. Roth (1 d, AMNH); Mendocino County: Fault Rock Cave, 2 January 1960, R.E. Graham (#1622, 2 juveniles; #1623, 2 $; #1625, 2 d, 2 $, AMNH); 4.2 miles S. Piercy, 39°57'N, 123°47'W, 17 February 1967, V. Roth (1 d, AMNH); Yolo County: 18.5 km ESE Lower Lake, 38°52'N, 122°23'W, 14. v-7-1993, B.L. Fisher, pitfall traps, non-serpentine, chaparral (1 d, CASC). Diagnosis. — Distinguished from other Cal- ileptoneta, except C. briggsi, C. califomica, and C. sylva, by males having a proximal bulb process (Figs. 38-40, 59-61), and a straight retroapical seta (Fig. 39); from C. briggsi and C. sylva by having the proximal bulb process (Figs. 59-61) elongate, process length 1.0- 2.04 X bulb width, and having the accessory lobe reduced (Fig. 64); from C. califomica by having an elongate palpal tibia, length 1.02- 1.19 X carapace width, whereby the proximal bulb process does not reach the base of the tibia, bulb length 0.81-1.04 X tibia length. Male (holotype). — Total length 2.29. Col- oration and markings same as for C. calif ar- nica, except considerably darker throughout. Carapace 1.13 long, 0.93 wide, height at fo- vea 0.39 X carapace width; clypeus 0.17 high, chelicerae 0.7 1 long, fang furrow with 5 teeth along a narrow ridge and 5 denticles on retro- margin (Fig. 21). Ocular area 0.30 long, 0.23 wide; diameter PME 0.75 X PLE interdist- 252 THE JOURNAL OF ARACHNOLOGY Figures 53-58. — Calileptoneta cokeudolpheri new species, male from H. J. Andrews, right palpus. 53. retrolateral. 54. ventral. 55. prolateral. 56. retroventral. 57. ventral. 58. proapical. AH = apical hook, E = embolus, PF = proapical flange, PL = prolateral lobe, PS = paraembolar setae. Scale bars: 53 = 100 jjim, 54 = 20 p.m, 55 = 100 fxm, 56-58 = 30 pm. ances. Sternum 0.67 long, 0.70 wide; labium 0.08 long, 0.15 wide; palpal coxae 0.55 long, 0.21 wide. Spination: palpus: femur p2-l-2-3-3-2-3-3- 2-3-3, rl-1-2-1-1-1-2-1-2-2-2-2-2-1-1, vl (apical); patella dl, tibia r2, tarsus rl. Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]): I: 2.16 + 0.34 + 1.90 + 1.60 T 1.00 = [7.00]; II: 1.74 + 0.32 + 1.77 + 1.43 + 0.95 = [6.21]; III: 1.47 -f 0.30 + 1.31 + 1.25 + 0.83 = [5.16]; IV: 1.86 + 0.30 -t- 1.86 + 1.60 + LOO = [6.62]; pedipalpus: 1.53 + 0.64 + 1.02 + 0.36 = [3.55]. Femur I 2.32 X carapace width, palpal femur 1.65 X carapace width. Palpal bulb (Figs. 59-64) 1.0 long, 0.25 LEDFORD— REVISION OF CALILEPTONETA 253 Figures 59— 64.— Calileptoneta helferi (Gertsch), male from Mt. Diablo, right palpus. 59. retrolateral. 60. ventral. 61. prolateral. 62. retrolateral. 63. ventral. 64. prolateral. AC= apical constriction, AH = apical hook, AL = apical lobe, E = embolus, PBP = proximal bulb process, PF = proapical flange, PL = prolateral lobe, PS = paraembolar setae, RS = retroapical setae. Scale bars; 59 = 300 |xm, 60 = 231 [xm, 61 = 300 |xm, 62 = 43 |xm, 63 = 75 |xm, 64 = 86 pm. wide; palpal tibia with a retroapical group of stiff setae; proximal bulb process elongate, not reaching to base of tibia, bulb length 0.98 X length tibia; embolus narrowly forked at apex; paraembolar setae circular, reaching to base of fork on embolus (Figs. 63, 64); prolateral api- cal lobe reduced (Fig. 64). Abdomen dusky with pale chevron pattern (Fig. 6), 1.16 long, 1.05 wide. Variation {n = 9). — Total length 2.04- 2.34; carapace length 1.15-1.29 X carapace width; OAL 1.14-1.45 X OAW, diameter PME 0.50-0.73 X PLE interdistances; length femur I 2.15-2.66 X carapace width, palpal femur 1.34-1.98 X carapace width; palpal bulb length 0.81-1.10 X palpal tibia length; proximal bulb process length 1.40-2.04 X bulb width. Female (Claremont Ave.). — -Total length 2.25. Coloration and markings same as male. Carapace 0.92 long, 0.73 wide, height at fo- vea 0.26 X carapace width; clypeus 0.12 high, chelicerae 0.45 long, fang furrow with 7 teeth on a narrow ridge and 3 denticles on retro- margin (Fig. 21). Ocular area 0.25 long, 0.22 wide; diameter PME 0.50 X PLE interdist- ances. Sternum 0.57 long, 0.51 wide; labium 0.08 long, 0.15 wide; palpal coxae 0.36 long, 0.18 wide. Spination: palpus: patella dl, tarsus p2-l-l, rl-1, vl-2 (apical). Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]): I: 1.47 + 0.30 + 1.62 + 0.95 + 0.87 - [5.21]; II: 1.23 4 0.28 + 1.11 + 0.97 + 0.71 - [4.30]; III: 1.03 + 0.28 + 0.95 + 0.85 + 0.61 = [3.72]; IV: 1.37 + 0.26 + 1.25 + 254 THE JOURNAL OF ARACHNOLOGY Figures 65-70. — Calileptoneta noyoana (Gertsch), male from Fort Bragg, right palpus. Arrow on 68 to retroapical setae. 65. retrolateral. 66. ventral. 67. prolateral. 68. retrolateral. 69. ventroapical. 70. retrola- teral. AC = apical constriction, AH = apical hook, AL = apical lobe, E = embolus, PF = proapical flange, PL == prolateral lobe, PS = paraembolar setae, RS = retroapical setae. Scale bars: 65-67 = 200 pm, 68-70 = 75 pm. 1.09 + 0.75 = [4.72]; pedipalpus: 0.54 + 0.17 + 0.39 + 0.49 = [1.59]. Femur I 2.01 X car- apace width, palpal femur 0.74 X carapace width. Abdomen 1.33 long, 1.06 wide. Atrium 0.14 long, 0.19 wide, spermathecae 0.13 long (Fig. 115). Variation {n = 3). — Total length 2.24- 2.40; carapace length 1.21-1.26 X carapace width; OAL 1.14-1.19 X OAW, diameter PME 0.50-0.64 X PLE interdistances; length femur I 2.01-2.13 X carapace width, palpal femur 0.74-0.80 X carapace width; atrium length 0.74-0.81 X width, spermathecae 0.68-1.18 X atrium width. Natural history. — The Mt. Diablo speci- mens were found under moist stones in oak woodland sympatrically with C. californica. Distribution. — Northwestern California (Fig. 120). Calileptoneta noyoana (Gertsch 1974) Figs. 4-5, 19, 65-70, 96, 100-101, 106-108, 120 Leptoneta noyoana Gertsch 1974: 196-197; Brig- noli 1977: 217. Calileptoneta noyoana (Gertsch): Platnick 1986: 15; Platnick 2002. Type material. — Male holotype from 12- 15 miles E. Noyo, Mendocino County, Cali- fornia, USA, 39°25'N, 123°48'W, 13 Septem- ber 1961, W.J. Gertsch, W. Ivie (AMNH, examined) Material examined. — USA: California: Humboldt County: Humboldt Redwoods State Park, Founder’s Grove, 40°21'N, 123°55'W, LEDFORD— REVISION OF CALILEPTONETA 255 Figures 71—76. — Calileptoneta oasa (Gertsch), male from Andreas Canyon, right palpus. 71. retrolateral. 72. ventral. 73. prolateral. 74. retroapical. 75. ventroapical. 76. proapical. AC = apical constriction, AH = apical hook, AL = apical lobe, E = embolus, PF = proapical flange, PL = prolateral lobe, PS — paraembolar setae, RS = retroapical setae. Scale bars: 71-73 = 136 p.m, 74-76 = 75 jxm. 28 October 1990, in redwood duff, D. Ubick, W. Rauscher (29,2 juveniles including pen- ultimate male, DU); Humboldt Redwoods State Park, Tall Tree, 40°19'N, 123°59'W, 250 ft. elevation, 11 September 1992, D. Ubick, J. Boutin (39,2 juveniles including penultimate male, DU); Mendocino County: Jackson State Forest, Camp Dunlap Area, 39°2UN, 123°33'W, 16 September 1990, in redwood duff, D. Ubick (Id, DU); 0.5 miles W. Camp Dunlap, 39°2rN, 123°33'W, 400 ft. elevation, 16 September 1990, in redwood duff, D. Ubick (1 9, DU); Dunlap Camp, 39°2UN, 123°33'W, 400 ft. elevation, 7 September 1992, in redwood duff, D. Ubick, J. Boutin (1 9, DU); 3.0 miles S. Rockport, 39°44'N, 123°48'W, 300 ft. elevation, 19 September 1990, in redwood duff, D. Ubick (1 d, DU); 2.0 miles S. Usal Campground, 39°50'N, 123°50'W, 1000 ft. elevation, 19 September 1990, in redwood duff, D. Ubick (1 9, DU); 1 miles NE Usal Road along HWY 1, 200 ft. elevation, 39°50'N, 123°50'W, 20 September 1990, D. Ubick (Id, DU); Big River Camp, ~2 miles W. James Creek, 39°20'N, 123°30'W, 5 May 1991, in redwood duff, D. Ubick (2 d, 2 9,3 juveniles, DU); Noyo Riv- er, 14.5 air miles E. Fort Bragg, 39°25.5'N, 123°32'W, 25-26.V.1996, C.E. Griswold (2 d, 1 9 , juvenile, CASC). Diagnosis. — Distinguished from other Cal- ileptoneta, except C. wapiti, by having males with retrolateral twisted tarsal setae on the palpus (Figs. 37, 68) and an enlarged distal cheliceral tooth (Fig. 19); from C. wapiti by having an elongate palpal femur, 2.24-2.78 X carapace width (Fig. 5). Male (holotype). — Total length 2.26. Spec- 256 THE JOURNAL OF ARACHNOLOGY Figures 77-82. — Calileptoneta sylva (Chamberlin & Ivie), male from Samwell Cave, right palpus. 77. retrolateral. 78. ventral. 79. prolateral. 80. retroapical. 81. ventroapical. 82. retroapical. AC = apical constriction, AH = apical hook, AL = apical lobe, E = embolus, PBP = proximal bulb process, PF = proapical flange, PL = prolateral lobe, PS = paraembolar setae, RS = retroapical setae. Scale bars: 77- 79 = 200 |jLm, 80-82 = 86 jim. imen faded. Carapace yellow-brown with dusky mottling along margin; clypeus yellow- brown; sternum yellow-brown; coxae, tro- chanters, legs, and pedipalpi with dusky mot- tling, being especially conspicuous at the bases and apices of the segments. Carapace 1.06 long, 0.87 wide, height at fo- vea 0.46 X carapace width; clypeus 0.15 high, chelicerae 0.74 long, fang furrow with 8 teeth along a narrow ridge and 5 denticles on retro- margin (Fig. 19). Ocular area 0.30 long, 0.22 wide; diameter PME 0.70 X PLE interdist- ances. Sternum 0.63 long, 0.58 wide; labium 0.10 long, 0.18 wide; palpal coxae 0.50 long, 0.24 wide. Spination: palpus: patella dl, tibia r3-l-l, tarsus pi (apical), rl-1-1 (apical). Leg mea- surements (Eemur + Patella + Tibia + Meta- tarsus -f Tarsus = [Total]): I: 2.04 + 0.34 + 2.17 + 1.70 + 1.02 = [7.27]; II: 1.56 + 0.32 + 1.60 + 1.34 + 0.93 = [5.74]; III: 1.29 + 0.24 + 1.24 + 1.15 + 0.73 - [4.65]; IV: 1.73 + 0.29 + 1.70 + 1.44 + 0.90 - [6.06]; pe- dipalpus: 2.42 + 1.31 + 1.21 + 0.46 = [5.40]. Femur I 2.34 X carapace width, palpal femur 2.78 X carapace width. Palpal bulb (Figs. 65-70) 0.49 long, 0.27 wide; retroapical seta proximally broad and tapering to a point distally; embolus broadly forked at apex; paraembolar setae fan-like, with 3 truncate setae at the base, and a single seta forming a broad flange distally; accessory lobe reduced (Fig. 70). Abdomen dusky with pale chevron pattern (Fig. 4), 1.20 long, 0.96 wide. Variation (n = 4). — Total length 2.17-2.50; carapace length 1.21-1.59 X carapace width; OAL 1.20-1.37 X OAW, diameter PME 0.60- LEDFORD— REVISION OF CALILEPTONETA 257 Figures 83-88. — Calileptoneta ubicki new species, male from Arroyo Seco Canyon, right palpus. 83. retrolateral. 84. ventral. 85. prolateral. 86. retrolateral. 87. ventroapical, 88. proapical. AC = apical con- striction, AH = apical hook, AL = apical lobe, E = embolus, PF = proapical flange, PL = prolateral lobe, PS = paraembolar setae, RS = retroapical setae. Scale bars: 83 = 250 ixm, 84 = 231 fxm, 85 = 270 fxm, 86 = 136 pm, 87 = 86 pm, 88 = 75 pm. 0.70 X PLE interdistances; length femur I 2.32-2.35 X carapace width, palpal femur 2.24-2.78 X carapace width; bulb length 0.40- 0.45 X palpal tibia length; abdomen pale to dusky, with or without chevron pattern. Female (Fort Bragg)* — Total length 2.24. Coloration and markings same as male. Car- apace 0.69 long, 0.59 wide, height at fovea 0.50 X carapace width; clypeus 0.11 high, chelicerae 0.33 long, fang furrow with 8 teeth along a narrow ridge and 3 denticles on retro- margin (Fig, 19). Ocular area 0.24 long, 0.19 wide; diameter PME 0.50 X PLE ieterdist- ances. Sternum 0.54 long, 0.56 wide; labium 0,09 long, 0.16 wide; palpal coxae 0.36 long, 0.20 wide. Spination: palpus: patella dl, tarsus p2-l-l (apical), rl-1 (apical). Leg Measurements (Fe- mur + Patella + Tibia + Metatarsus + Tarsus - [Total]): I: 1.41 A 0.29 + 1.54 A 1.15 A 0.85 - [5.24]; II: 1.17 A 0.29 A 1.20 A 0.93 A 0.73 - [4.32]; III: 1.04 A 0.24 A 0.90 A 0.83 A 0.61 = [4.19]; IV: 1.37 A 0.24 A 1.61 A 1,36 A 0.88 = [5.46]; pedipalpus: 0.58 A 0.23 A 0.45 A 0.50 = [1.76]. Femur I 1.83 X carapace width, palpal femur 0.75 X carapace width. Abdomen 1.31 long, 0.96 wide. Atrium 0.19 long, 0.24 wide, spermathecae 0.17 long (Figs. 106-108). Variation (w ~ 4). — Total length 2.03- 2.53; carapace length L0-L23 X carapace width; OAL 1.0-1.35 X OAW, diameter PME 0.6-0.7 X PLE interdistances; length femur I 1.62-2.08 X carapace width, palpal femur 0.75-0.83 X carapace width; atrium length 0.77-0.79 X width, spermathecae 0.65-0.71 X atrium width. 258 THE JOURNAL OF ARACHNOLOGY Figures 89-94. — Calileptoneta wapiti (Gertsch), male from Cameron Road, right palpus. 89. retrolateral. 90. ventral. 91. prolateral. 92. retroapical. 93. ventroapical. 94. proapical. AC= apical constriction, AH = apical hook, AL = apical lobe, E = embolus, PF = proapical flange, PL = prolateral lobe, PS = paraembolar setae, RS == retroapical setae. Scale bars: 89-91 = 176 p,m, 92-94 = 86 p.m. Natural history, — This spider is most commonly found among the dense leaf litter in redwood and Douglas hr forests although they may also be found under moist rotting logs. Individuals in captivity constructed small sheet webs like those of other Calilep- toneta species. The extremely long palp of the male suggests a mating behavior potentially unique among Calileptoneta species. Distribution. — Restricted to redwood {Se- quoia sempervirens) and mixed evergreen for- ests (Douglas hr-redwood) on the Pacihc northwest coast of California (Fig. 120). Calileptoneta oasa (Gertsch 1974) Figs. 15, 32-34, 71-76, 102-103, 119 Leptoneta oasa Gertsch 1974: 197; Brignoli 1977: 217. Calileptoneta oasa (Gertsch): Platnick 1986: 15; Platnick 2002. Type material. — Male holotype from An- dreas Canyon, off Palm Canyon, near Palm Springs, Riverside County, California, USA, 33°49'N, 116°32'W, 26 March 1960, W.J. Gertsch (AMNH, examined). Paratype: USA: California: 1 9, same data as holotype (AMNH, examined). Other material examined. — USA: Cali- fornia: Riverside County: Andreas Canyon, near Palm Springs, 33°49'N, 116°32'W, 3 March 1956, V. Roth (1 d, 1 9, 1 juvenile, MCZ unique # 35493). Diagnosis. — Distinguished from other Cal- ileptoneta, except C. ubicki, by having fe- males with a bihd atrium (Fig. 102) and males without a proximal bulb process (Figs. 32-34) or retrolateral twisted tarsal setae on the pal- pus (Fig. 37); from C. ubicki by the following combination of characters: lacking a retrodis- tal cheliceral apophysis (Fig. 16), having a hook-shaped retroapical tibial seta on the pal- LEDFORD— REVISION OF CALILEPTONETA 259 Figures 95-97. — Calileptoneta sp., tarsal organs, right palpus, dorsoapical. 95. C ubicki new species, from Arroyo Seco Canyon. 96. C. noyoana (Gertsch), from Fort Bragg. 97. C. sylva (Chamberlin & Ivie), from Samwell Cave. Scale bars: 95 = 7.5 pm, 96 = 10 pm, 97 = 8.6 pm. pus (Fig. 71), and having the proapical flange straight anteriorally (Fig. 76). Male (holotype). — Total length 1 .70. Spec- imen faded. Carapace yellow-brown with dusky mottling surrounding margin, and lat- erally along caput margins; clypeus with dusky mottling distally; sternum dusky; cox- ae, trochanters, leg segments, and palpi yel- low-brown. Carapace 0.75 long, 0.62 wide, height at fo- vea 0.54 X carapace width; clypeus 0.12 high, chelicerae 0.45 long, fang furrow with 5 teeth along a narrow ridge and 1 denticles on retro- margin (Fig. 15). Ocular area 0.21 long, 0.15 wide; diameter PME 0.75 X PLE interdist- ances. Sternum 0.48 long, 0.45 wide; labium 0.08 long, 0.15 wide; palpal coxae 0.30 long, 0.15 wide. Spination: palpus: patella dl; tibia rl-L Leg measurements (Femur + Patella + Tibia + 260 THE JOURNAL OF ARACHNOLOGY Figures 98-101. — Calileptoneta sp., female genitalia, ventral. 98. C. ubicki new species, from Arroyo Seco Canyon. 99. C. colifornicci (Banks), from Bell Station. 100. C. noyoana (Gertsch), from Fort Bragg. 101. C. noyoana (Gertsch), from Fort Bragg, left lateral, arrow to shaip bend in spermathecae. AB = apical bifurcation, S = spermathecae. Scale bars: 98 = 0.08 mm, 99 = 0.10 mm, 100 = 0.12 mm, 101 = 0.08 mm. Illustrations by JL. Metatarsus + Tarsus = [Total]): I: 1.83 + 0.24 + 1.85 + 1.65 + 0.98 = [6.55]; II: 1.37 + 0.24 + 1.51 T missing + missing = [N/A]; III: 1.12 + 0.20 + 1.05 + 1.0 + 0.61 = [3.98]; IV: 1.46 + 0.20 + 1.07 + 1.0 + 0.61 = [4.34]; pedipalpus: 0.42 + 0.15 + 0.20 + 0.29 = [1.06]. Femur I 2.95 X carapace width, palpal femur 0.68 X carapace width. Palpal bulb (Figs. 71-76) 0,36 long, 0.20 wide; palpal tibia with a retroapical hook- shaped seta; embolus tapering to a sharp point (Fig. 75); paraembolar setae fan-like with 3 sagitate setae extending to apex of embolus (Fig. 76); accessory lobe reduced. Abdomen dusky with pale chevron pattern, 0.95 long, 0.77 wide. Variation {n = 2). — Total length 1.70- 1.71; carapace length 1.21-1.23 X carapace width; OAL 1.08-1.40 X OAW, diameter PME 0.66-0.77 X PLE interdistances; length femur I 2.79-3.0 X carapace width, palpal fe- mur 0.68-0.69 X carapace width; bulb length 1.80-2.20 X palpal tibia length. Female (paratype). — Total length 1.28. Specimen faded. Coloration and markings same as male. Carapace 0.69 long, 0.59 wide, height at fo- vea 0.50 X carapace width; clypeus 0. 1 1 high, chelicerae 0.33 long, fang furrow with 6 teeth on a narrow ridge and 4 denticles on retro- margin (Eig. 15). Ocular area 0.18 long, 0.14 wide; diameter PME 0.62 X PLE interdist- ances. Sternum 0.45 long, 0.42 wide; labium 0.05 long, 0.15 wide; palpal coxae 0.25 long, 0. 13 wide. Spination: palpus: patella dl, tarsus pi, rl- 1, v2-L Leg measurements (Eemur + Patella + Tibia T Metatarsus + Tarsus = [Total]): I: 1.29 + 0.22 + 1.39 + 1.12 + 0.78 = [4.8]; II: 1.0 + 0.20 + 1.0 + 0.83 + 0.61 = [3.64]; III: 0.89 + 0.20 + 0.78 T 0.71 + 0.51 = [3.09]; IV: 1.17 + 0.20 + 1.02 + 0.90 + 0.49 = [3.78]; pedipalpus: 0.41 + 0.12 + 0.22 + 0.38 = [1.13]. Eemur I 2.19 X carapace width, palpal femur 0.69 X carapace width. Abdomen 1.19 long, 0.95 wide. Atrium LEDFORD— REVISION OF CALILEPTONETA 261 Figures 102-105. — Calileptoneta sp., female genitalia, ventral (102, 104) and left lateral (103, 105). 102-103. C. oasa (Gertsch), from Andreas Canyon. 104-105. C. ubicki new species, from Arroyo Seco Canyon. AB = apical bifurcation, S = spermatheca. Scale bars: 102 = 0.10 mm, 103 = 0.05 mm, 104 = 0.10 mm, 105 = 0.08 mm. 0.18 long, 0.21 wide, spermathecae 0.12 long (Figs. 102-103). Variation (#i = 2). — Total length 1.28- 1.72; carapace length 1.17-1.22 X carapace width; OAL 1.24-1.29 X OAW, diameter PME 0.62-0.70 X PLE interdistances; length femur I 2.19-2.56 X carapace width, palpal femur 0.69-0.72 X carapace width. Natural history. — Unknown. Distribution. — Known only from the type locality (Fig, 119). Calileptoneta sylva (Chamberlin & Ivie 1942) Figs. 23, 77-82, 97, 109-110, 121 Leptoneta sylva Chamberlin & Ivie 1942: 9-10. Leptoneta californica (Chamberlin & Ivie): Gertsch 1974: 191 (misidentification, not L. californica Banks 1904). Calileptoneta californica (Chamberlin & Ivie) (part): Platnick 1986: 14; Platnick 2002. Type material. — Female holotype, 19 miles N. Wolf Creek, Jackson County, Oregon, USA, 42°42'N, 122°57'W, 6 April 1937, J.C. Chamberlin (holotype female, AMNH). Other material examined. — USA: Cali- fornia: Shasta County: Samwell Cave, 40°55'N, 122°14'W, 14 April 2000, J.M. Led- ford, at entrance under limestone (1 d, 2 ju- veniles, CASC); Oregon: Jackson County: Ashland watershed, T39S, RIE, SEC 34, PFT #46, 20 July 1998, R.W. Peck et al. (1 d, OSU); Jenny Creek LSR, Medford district, 41°58'N, 122°24'W, BLM, LS, Oldgrowth, T39S, R03E, SEC 35, PFT 31-40, 21-23 June 262 THE JOURNAL OF ARACHNOLOGY I 106 ■| A Figures 106-108. — Calileptcmeta noyoana (Gertsch), female genitalia, ventral (106) and left lateral (107, 108). Arrow in 107 to sharp bend in spermatheca. Scale bars: 106 = 0.10 mm, 107 = 0.05 mm, 108 = 0.025 mm. 1999, B. Peck (1 S , OSU); Jenny Creek LSR, Medford district, 41°58'N, 122°24'W, BUM, LS, Oldgrowth, T39S, R03E, SEC 35, traps 21-3-0, 16-18 August 1999, B. Peck (1 $, OSU). Diagnosis. — Distinguished from other Cal- ileptoneta, except C. briggsi, C. californica, and C. helferi, by males having a proximal bulb process (Pigs. 38-40, 77-79) and a straight retroapical seta (Figs. 39, 78); from C. californica and C helferi by having the proximal bulb process (Figs. 77-79) short, process length 0.55-0.86 X bulb width, and having the prolateral apical lobe large (Figs. 46, 82); from C. briggsi by being darkly pig- mented and having a straight, distally nar- rowed embolus with a slight prolateral bend (Fig. 81). Male (Samwell Cave). — Total length 2.17. Carapace yellow-brown with fine dusky mot- tling surrounding margin, and laterally along caput margins; clypeus with dusky mottling distally; sternum dusky; coxae, trochanters, legs, and pedipalpi with dusky mottling, being especially conspicuous at the bases and apices of the segments. LEDFORD— REVISION OF CALILEPTONETA 263 Figures 109—112. — Calileptoneta sp., female genitalia, ventral. 109. C. sylva (Chamberlin & Ivie) ho- lotype. 110. C. sylva (Chamberlin & Ivie), from Jenny Creek. 111-112. C briggsi new species from Indian Valley Creek Cave. Scale bars: 109-1 10 = 0.10 mm, 1 1 1-1 12 = 0.12 mm. Carapace 1.0 long, 0,90 wide, height at fo- vea 0.24 X carapace width; clypeus 0.14 high, chelicerae 0.57 long, fang furrow with 9 teeth along a narrow ridge and 4 denticles on retro- margin (Fig. 23). Ocular area 0.26 long, 0.22 wide; diameter PME 0.64 X PLE interdist- ances. Sternum 0.65 long, 0.62 wide; labium 0.10 long, 0.19 wide; palpal coxae 0.47 long, 0.21 wide. Spination: palpus: femur p 1-3-2- 1-2- 1-2-2- 2, r5-3-2-2 (apical), vl (apical); patella dl; tibia rl -2- 1-1-3; tarsus rl (apical). Leg mea- surements (Femur + Patella + Tibia + Meta- tarsus + Tarsus ^ [Total]): I: 2.36 + 0.34 + 2.72 + 2.18 + 1.15 = [8.75]; II: 1.74 + 0.32 + 1.74 + 1.47 + 0.88 = [6.15]; III: 1.43 + 0.28 + 1.29 + I.2I + 0.73 = [4.94]; IV: I.8I + 0.30 + 1.77 + 1.61 + 0.95 = [6.44]; pe- dipalpus: 0.95 + 0.36 + 0.49 + 0.63 = [2.43]. Femur I 2.62 X carapace width, palpal femur 0.1.05 X carapace width. Palpal bulb (Figs. 77-82) 0.61 long, 0.22 wide; palpal tibia with a retroapical group of stiff setae; proximal bulb process short, reach- ing to base of tibia, bulb length 1.24 X length tibia; embolus distally narrowed with a slight prolateral bend; paraembolar setae circular, distally broad, reaching to apex of embolus (Fig. 82). Abdomen dusky with pale chevron pattern, 1.17 long, 0.89 wide. Variation {n — 2). — Total length 2.04- 2.17; OAL 1.18-1.23 X OAW, diameter PME 0.50-0.64 X PLE interdistances; length femur I 2.16-2.72 X carapace width, palpal femur 0.96-1.05 X carapace width; palpal bulb length 1.20-1.24 X palpal tibia length; prox- imal bulb process length 0.55-0.86 X bulb width. Female (holotype). — Total length 2.69. Specimen faded. Carapace yellow-brown with dusky mottling surrounding margin, and lat- 264 THE JOURNAL OF ARACHNOLOGY 113 114 Figures 113—116. — Calileptoneta sp., female genitalia, ventral. 113. C. califomica (Banks) from Oak- ville. 114. C. califomica (Banks) from Bell Station. 115. C. helferi (Gertsch) from Fault Rock Cave. 116. C. helferi (Gertsch) from Claremont Avenue. Scale bar = 0.10 mm. erally along caput margins; clypeus with dusky mottling distally; sternum yellow- brown; coxae, trochanters, leg segments, and palpi yellow-brown. Carapace 1 .05 long, 0.92 wide, height at fo- vea 0.26 X carapace width; clypeus 0.10 high, chelicerae 0.61 long, fang furrow with 9 teeth on a narrow ridge and 2 denticles on retro- margin (Fig. 23). Ocular area 0.28 long, 0.23 wide; diameter PME 0.54 X PLE interdist- ances. Sternum 0.65 long, 0.59 wide; labium 0.10 long, 0.16 wide; palpal coxae 0.45 long, 0.23 wide. Spination: palpus: patella dl, tarsus p2-l-l, rl-1, vl-2 (apical). Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]): I: 1.77 + 0.34 + 2.0 + 1.61 + 1.07 = [6.79]; II: 1. 41 + 0.34 + missing + missing + missing = [N/A]; III: 1.29 + 0.28 + 1.21 + 1.11 + 0.75 = [4.64]; IV: 1.70 + 0.32 + 1.70 + 1.47 + 0.91 = [6.10]; pedipalpus: 0.66 + 0.23 + 0.47 + 0.59 = [1.95]. Femur I 1.92 X carapace width, palpal femur 0.72 X cara- pace width. Abdomen 1.64 long, 1.53 wide. Atrium 0.19 long, 0.20 wide, spermathecae 0.16 long (Fig. 109). Variation {n = 2). — The only known fe- male besides the holotype (Jenny Creek) is badly damaged and missing most appendages. Atrium 0.19 long, 0.20 wide, spermathecae 0. 14 long (Fig. 1 10). Remarks. — Gertsch (1974) synonymized L. sylva with L. califomica based on the sim- ilarity in abdominal pattern and female geni- talia between the type of L. sylva and a to- potypic female of L. califomica (Mt. Diablo). The type of L. califomica was lost at the Cal- ifornia Academy of Sciences during the 1906 earthquake and fire. Since the abdominal pat- terns within Calileptoneta species are vari- able, and the female genitalia for species out- side the oasa group do not allow specific diagnosis, I consider the synonymy of these LEDFORD— REVISION OF CALILEPTONETA 265 Figures 117-118. — Calileptoneta cokendolpheri new species, female genitalia, ventral. 1 17-118. Fe- males from H. J. Andrews, showing intraspecific variation. Scale bar = 0.12 mm. species unjustified. Additionally, males be- longing to the califomica group, clearly dif- ferent from the species treated in this paper, have been recently discovered near the type locality of L. sylva. Instead of assigning a new name to this species, they are placed with L. sylva. Natural history. — The male from Samwell Cave, Shasta County was collected under limestone at the cave entrance that remains cool and moist throughout the year. No spec- imens have ever been taken inside the cave despite extensive arachnid surveys. Distribution.- — Southern Oregon to north- central California (Fig. 121). Calileptoneta uMcki new species Figs. 1, 2, 3, 16, 83-88, 95, 98, 104, 105, 119 Type material. — Male holotype from Ar- royo Seco Canyon Campground, SW of Lakes, Monterey County, California, USA, 36°14'N, 12r28'W, 22 January 2001, J.M. Ledford, P. Marek (CASC). Other material examined. — USA: Cali- fornia: Monterey County: Arroyo Seco Camp- ground, SW of Lakes, 36°14'N, 12r28'W, Figure 119. — Distribution map for Calileptoneta oasa (Gertsch), and Calileptoneta ubicki new spe- cies • = C. oasa, ■ = C. ubicki. 900 ft. elevation, under granite, 6 May 1995, D. Ubick, W. Savary (1 S , DU), 22 January 2001, J.M. Ledford, P. Marek (2 $, CASC). Etymology.— This species is named in honor of Mr. Darrell Ubick, collector of this and many other leptonetid spiders throughout California. Diagnosis. — Distinguished from other Cal- ileptoneta, except C. oasa, by having females with a bifid atrium (Figs. 98, 104), and males without a proximal bulb process (Figs. 32-34, 83-88) or retrolateral twisted tarsal setae on the palpus (Fig. 37); from C oasa by males having a retrodistal cheliceral apophysis (Fig. 16), a whip-shaped retroapical tibial seta on the palpus (Fig. 86), and having a sinuate proapical flange (Fig. 88). Male (holotype). — Total length 2.31. Car- apace yellow-brown with a fine dusky band surrounding margin, and laterally along caput margins; clypeus with faint dusky mottling distally; sternum dusky; coxae, trochanters, legs, and pedipalpi with dusky mottling, being especially conspicuous at the bases and apices of the segments. Carapace 1.08 long, 0.91 wide, height at fo- vea 0.44 X carapace width; clypeus 0.19 high, chelicerae 0.93 long, fang furrow with 7 teeth along a narrow ridge and 5 denticles and a large distal tooth on retromargin (Fig, 16). Oc- 266 THE JOURNAL OF ARACHNOLOGY Figure 120. — Distribution map for Calileptoneta sp. ■ = C. californica (Banks), • = C. helferi (Gertsch), o = C. californica incertae sedis, A = C. noyoana (Gertsch), A = C. wapiti (Gertsch). Figure 121. — Distribution map for Calileptoneta briggsi new species, Calileptoneta cokendolpheri new species, and Calileptoneta sylva (Chamberlin & Ivie). A — Calileptoneta briggsi, ■ = C. cok- endolpheri, • = C. sylva. ular area 0.23 long, 0.22 wide; diameter PME 0.53 X PLE interdistances (Fig. 2). Sternum 0.67 long, 0.63 wide; labium 0.12 long, 0.19 wide; palpal coxae 0.53 long, 0.17 wide. Spination: palpus: patella dl, tibia rl-1-1. Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]): I: 2.41 + 0.34 T 2.80 + 2.43 -h 1.29 = [9.27]; II: 1.80 + 0.29 + 1.95 + 1.70 + 0.98 = [6.72]; III: 1.54 + 0.24 + 1.44 + 1.41 + 0.85 = [5.48]; IV: 2.0 + 0.32 + 1.90 + 1.80 + 1.0 = [7.02]; pedipalpus: 0.71 + 0.21 -f 0.38 + 0.36 = [1.66]. Femur I 1.09 X carapace width, palpal femur 0.78 X carapace width. LEDFORD— REVISION OF CALILEPTONETA 267 Figure 122. — Distribution map for undiagnosable females and juvenile Calileptoneta sp. A= undiag- Eosable females, • = juveniles. Palpal bulb (Figs. 83-88) 0.42 long, 0.25 wide; palpal tibia with a retroapical whip- shaped seta; embolus tapering to a sharp point; paraembolar setae fan-like with 3 da- vate setae (Fig. 88); proapical flange sinuate (Fig. 88); accessory lobe reduced. Abdomen pale with a dusky chevron pat- tern (Fig. 2), 1.23 long, 1.07 wide. Variation {m = 2). — Total length 2.30- 2.31; carapace length 1.17-1.19 X carapace width; OAL 1.04-1.05 X OAW, diameter PME 0.50-0.53 X PLE interdistances; length femur I 2.65-2.67 X carapace width, palpal femur 0.75-0.78 X carapace width; bulb length 1.30-1.56 X palpal tibia length. Female (paratype), — Total length 2.31. Coloration and markings same as male. Carapace 0.80 long, 0.68 wide, height at fo- vea 0.41 X carapace width; ciypeus 0.12 high, chelicerae 0.31 long, fang furrow with 8 teeth along retromargin and 4 denticles (Fig. 16). Ocular area 0.18 long, 0.19 wide; diameter PME 0.50 X PLE interdistances. Sternum 0.50 long, 0.47 wide; labium 0.06 long, 0.14 wide; palpal coxae 0.28 long, 0.13 wide. Spination: palpus: patella dl, tarsus pi -1-1, rl-l-l-v2 (apical). Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]): I: 1.60 + 0.27 + 1.70 + 1.46 + 0.98 = [6.01]; II: 1.24 + 0.22 + 1.21 + 1.07 + 0.78 = [4.52]; III: 1.07 + 0.22 + 0.93 + 0.90 + 0.83 = [3.95]; IV: 1.4 + 0.22 + 1.24 + 1.15 + 0.80 - [4.81]; pedipalpus: 0.42 + 0.14 + 0.29 + 0.50 - [1.35]. Femur I 2.4 X car- apace width, palpal femur 0.62 X carapace width. Abdomen 1.51 long, 1.13 wide. Atrium 0.19 long, 0.17 wide, spermathecae 0.14 long (Fig. 104). Variation (n = 2). — -Total length 2.31- 2.40; carapace length 1.22-1.18 X carapace width; OAL 0.95-1.0 X OAW, diameter PME 0.50 X PLE interdistances; length femur I 2.24-2.47 X carapace width, palpal femur 0.62-0.71 X carapace width; atrium length 1.0-1.12 X width, spermathecae 0.75-0.82 X atrium width. Natural history. — These spiders were found under moist rocks on a loose granitic slope. Individuals hung beneath tightly woven sheet webs 3-4cm in diameter (Fig. 1). Distribution. — Known only from the type locality (Fig. 119). Calileptoneta wapiti (Gertsch 1974) Figs. 18, 35-37, 89-94, 120 Leptoneta wapiti Gertsch 1974: 195; Brignoli 1977: 217. Calileptoneta wapiti (Gertsch): Platnick 1986: 15; Platnick 2002. Type material. — Male holotype from Cameron Road, near Elk, Mendocino County, California, USA, 39°07'N, 123°43'W, 16 Feb- ruary 1967, V. Roth (AMNH, examined). Other material examined. — USA: Cali- fornia: Mendocino County: Mendocino, 39°18'N, 123°47'W, 4 January 1958, L P. Hei- fer (1 (3, AMNH). Diagnosis. — Distinguished from other Cal- ileptoneta, except C. noyoana, by having males with retrolateral twisted tarsal setae on 268 THE JOURNAL OF ARACHNOLOGY the palpus (Figs. 37, 92) and an enlarged dis- tal cheliceral tooth (Fig. 18); from C. noyoana by having a short palpal femur, 0.63-1.0 X carapace width. Male (holotype)* — Total length 1.91. Spec- imen faded. Carapace and all appendages yel- low-brown. Carapace 1.31 long, 0.79 wide, height at fo- vea 0.38 X carapace width; clypeus 0.19 high, chelicerae 0.63 long, fang furrow with 7 teeth on a narrow ridge and 3 denticles on retro- margin (Fig. 18). Ocular area 0.22 long, 0.18 wide; diameter PME 0.05 X PLE interdist- ances. Sternum 0.59 long, 0.57 wide; labium 0.10 long, 0.15 wide; palpal coxae 0.46 long, 0.19 wide. Spination: palpus: patella dl; tibia rl-1. Leg measurements (Femur + Patella + Tibia + Metatarsus + Tarsus = [Total]): I: 1.78 + 0.29 + 1.90 + 1.49 + 0.98 = [6.44]; II: 1.80 + 0.29 + 1.44 T 1.20 T 0.80 = [5.53]; III: 1.15 + 0.27 + 1.07 + 1.02 + 0.73 = [4.24]; IV: 1.49 + 0.24 + 1.46 + 1.29 + 0.85 = [5.33]; pedipalpus: 0.78 + 0.29 + 0.42 + 0.36 = [1.85]. Femur I 2.25 X carapace width, palpal femur 1.0 X carapace width. Palpal bulb (Figs. 89-94) 0.45 long, 0.27 wide; retroapical seta proximally broad and tapering to a point distally; embolus broadly forked at apex; paraembolar setae circular, reaching to base of fork on embolus; acces- sory lobe small (Fig. 94). Abdomen dark, without chevron pattern, 1.16 long, 0.91 wide. Variation {n = 2). — Total length 1.91; car- apace length 1.20-1.66 X carapace width; OAL 1.22-1.36 X OAW, diameter PME 0.50- 0.75 X PLE interdistances; length femur I 2.09-2.25 X carapace width, palpal femur 0.67-1.0 X carapace width, bulb length 1,0— 1.07 X palpal tibia length. F emale. — U nkno wn . Natural history. — Trips to relocate this rare species have proven unsuccessful. Spec- imens originally determined as C wapiti by Gertsch (1974) were females and juveniles with no diagnostic features. Furthermore, these specimens fit into the geographical range of C helferi and C. noyoana (Fig. 120). Due to the difficulty in the determination of females and the sympatric distribution of these species in Mendocino County, no female specimens are currently assigned to C. wapiti. Distribution. — Mendocino County, North- ern California (Fig. 120), ACKNOWLEDGMENTS Principal support for this project came from the Exline-Frizzell Fund and Entomology De- partment at the California Academy of Sci- ences. Research at Sequoia and King's Can- yon National Park was made possible through a research permit from the National Park Ser- vice, facilitated by Dr. David Graber. Research at Mt. Diablo State Park and Humboldt Red- woods State Park was made possible through a research permit from the California State Park Service. Additional fieldwork was a central compo- nent of this study and I wish to thank Mr. Paul Marek (CAS), Dr. Tom Briggs (CAS), Mr. Joel Despain (NPS), Ms. Karin Ledford, and especially Mr. Dairell Ubick (CAS) and Ms. Suzanne Ubick (CAS) for assistance in the field. All habitus illustrations are by Ms. Vir- ginia Kirsch. Mr. Darrell Ubick (CAS) assist- ed with the scanning electron microscope. Mr. James Cokendolpher, Mr. Darrell Ubick (CAS) and Dr. Norman Platnick (AMNH) provided suggestions and helped interpret the oftentimes difficult morphology of these spi- ders. Dr. Charles Griswold (CAS), Dr. Greg Spicer (SFSU) and Dr. Bob Patterson (SFSU), served as advisors on my thesis committee and assisted in ways too numerous to mention. Finally, I would like to thank Dr. Charles Griswold, Ms. Karin Ledford, Mr. Darrell Ubick, Ms. Suzanne Ubick and the students and staff at the California Academy of Sci- ences for encouragement and, without whom, this study would not have been possible. I would like to dedicate this study to the memory of Dr. Willis Gertsch, discoverer of many remarkable spider taxa throughout North America, and an inspiration to genera- tions of American arachnologists. A draft of this manuscript was critically read by James Cokendolpher, Charles Gris- wold, Bob Patterson, Greg Spicer and Dan-ell Ubick. LITERATURE CITED Banks, N. 1904. Some Arachnida from California. Proceedings of the California Academy of Sci- ences (3*'^^ ser.) 3:331-337. Banks, N. 1910. Catalogue of nearctic spiders. U.S, National Museum Bulletin 72:1-80. Brignoli, P.M. 1972. Some cavernicolus spiders LEDFORD— REVISION OF CALILEPTONETA 269 from Mexico (Araneae), in “Subterranean fauna of Mexico, Part I”. Quaderna Accademia Na- zionale dei Lincei 171:129-155. Brignoii, P.M. 1974. Notes on spiders, mainly cave- dwelling, of Southern Mexico and Guatemala. Quaderna Accademia Nazionale dei Lincei 171(2):195-238. Brignoii, P.M. 1975. Ueber die Gruppe der Haplo- gynae. Proceedings of the 6* International Ar- achnology Congress (Amsterdam 1974), pp. 33- 38. Brignoii, P.M. 1976. Ragni di Grecia IX. Specie nu- ove o interessanti delle famiglie Leptonetidae, Dysderidae, Pholcidae ed Agelenidae. Revue Suisse de Zoologie 83:539-578. Brignoii, P.M, 1977. Spiders from Mexico, III. A new leptonetid from Oaxaca, in “Subterranean fauna of Mexico, Part III”. Quaderna Accademia Nazionale dei Lincei 171:213-218. Brignoii, P.M. 1979a. On some cave spiders from Guatemala and United States (Araneae). Revue Suisse de Zoologie 86:435-443. Brignoii, P.M. 1979b. Ragni d’ Italia XXXL Specie cavernicole nuove o interessanti. Quaderni Per- iodico del Museo di Speleologica “V. Rivera”, Anno V N° 10 Dicembre 1979. Brignoii, P.M. 1979c. The morphology and the re- lationships of the Leptonetidae (Arachnida, Ar- aneae), Journal of Arachnology 7:231-236. Chamberlin, R.V. & W. Ivie. 1942. A hundred new species of American spiders. Bulletin of the Uni- versity of Utah, Biological Series 7:1-117. Coddington, J.A. 1983. A temporary slide mount allowing precise manipulation of small struc- tures. Verhandlungeii des Naturwissenschaftlich- en Vereins In Hamburg 26:291-292. Coddington, J.A. 1989. Spinneret silk morphology: evidence for the monophyly of orb-weaving spi- ders, Cyrtophorinae (Araneae), and the group Theridiidae plus Nesticidae. Journal of Arach- nology 17:71-95. Comstock J.H. 1913. The Spider Book. 721 pp. Cornell University Press, Ithaca, NY. Fage, L. 1913. Etudes sur les araignees cavernico- les, 11. Revision des Leptonetidae. Biospeologi- ca, Archives de Zoologie Experimentale et Ge- nerale (5 serie) 10:479-576. Gertsch, W.J. 1935. Spiders of the southwestern United States with descriptions of new species. American Museum Novitates 792:1-31. Gertsch, W.J. 1974. The spider family Leptonetidae in North America. Journal of Arachnology 1: 145-203. Machado, A. De Banos. 1941. Araignees nouvelles pour la faune portugaise 11. Pub, Inst. ZooL “Au- gust© Nobre”. Fac. Cienc. do Porto 3:1-60. Machado, A. De Barros. 1945. A propos de Fappareil respiratoir des “Leptonetidae” (Ara- neae). Pub. do Mus. ZooL do Porto 23:131-135. Moles, M.L. 1921. A list of California Arachnida: VII. Araneida or True Spiders. Journal of Ento- mology and Zoology, Claremont 13:39-45. Platnick, N.L 1986. On the tibial and patellar glands, relationships, and American genera of the spider family Leptonetidae (Arachnida, Ara- neae). American Museum Novitates 2855:1-16. Platnick, N.L 2002. The world spider catalog. Maintained by the American Museum of Natural History at: http://research.amnh.org/entomology/ spiders. Platnick, N.L, Coddington, J.A,, Forster, R.R., Gris- wold, C.E. 1991. Spinneret morphology and the phylogeny of haplogyne spiders. American Mu- seum Novitates 3016:1-73. Manuscript received 31 July 2002, revised 15 July 2003. 2004. The Journal of Arachnology 32:270-275 THREE NEW SPECIES OF THE SPIDER GENUS PHRUROLITHUS FROM CHINA (ARANEAE, CORINNIDAE) Chaeg-Min Yin: College of Life Science, Hunan Normal University, Changsha, Hunan Province 410081, PR. China Darrell Ubick: California Academy of Sciences, San Francisco, California 94118, USA You-Hui Bao and Xiang Xu: College of Life Science, Hunan Normal University, Changsha, Hunan Province 410081, PR. China ABSTRACT, Three new species of the genus Phrurolithus are described from the Gaoligong Mountain Region of Yunnan Province, China: Phrurolithus bifidus, P. qiqiensis and P. revolutus. Keywords: Taxonomy, Asia, Yunnan, Gaoligong Mountains The genus Phrurolithus C.L. Koch 1839, with some 70 described species, is currently the largest phrurolithine genus in the northern hemisphere (Platnick 2002). This richness, however, is deceptive as most of the species are probably not closely related to the type species, P. festivus (C.L. Koch 1835). For in- stance, the New World species currently in Phrurolithus, and representing over half of the taxa, are most probably all misplaced; some have already been formally reassigned to Sco- tinella Banks (Dondale & Redner 1982) and the others will no doubt follow. Additionally, some of the European species have recently been removed to Liophrurillus Wunderlich 1992 and Phrurolinillus Wunderlich 1995 (Wunderlich 1992, 1995) and some Asian ones to Otacilia Thorell 1 897 (Deeleman- Reinhold 2001) and what remains is probably still far from being monophyletic. For exam- ple, the 14 species currently recorded from China exhibit a wide spectrum of genitalic forms (Song, Zhu & Chen 1999). Males range in having the palpal tibial apophysis repre- sented by a single large process, similar to that in R. festivus (see figs. 240I-J in Song, Zhu & Chen 1999), to being polyfurcate (R. liaon- ingensis Song, Zhu & Chen 1999; see figs. 240M-N, in Song, Zhu & Chen 1999), to hav- ing one strongly reduced process (R. splendP dus Song & Zheng 1992; see figs. 241C-D, in Song, Zhu & Chen 1999), to completely lacking a process (R. daoxianensis Yin, Peng, Gong & Kim. 1997; see figs. 240G-H, Song, Zhu & Chen 1999). Similarly, and as would be expected, the female genitalia are equally diverse, with extreme variation in spermathe- cal size, the form of accessory bursae, and length and coiling of copulatory ducts. Given this morphological diversity, it is with some hesitation that we are placing the three new species in Phrurolithus. Whereas the three species do all have a pair of large spermathe- cae, as in R. festivus, the tibial apophysis of R. hifidus, the only male here described, is bifurcate and deviates significantly from that of the type species. Although this degree of intrageneric genitalic variation was accepted for Otacilia by Deeleman-Reinhold (2001), it remains to be seen if future studies will reveal a similar trend in Phrurolithus. The new species here described were col- lected in the Gaoligong Mountains by the first and second Sino-American expeditions. The type specimens are deposited in the College of Life Science at the Hunan Normal Univer- sity and some paratypes at the California Academy of Sciences. This is Scientific Con- tribution no. 27 from the California Academy of Sciences Center for Biodiversity Research and Information and contribution no. 20 from the China Natural History Project. METHODS Specimens were killed in 75% ethanol and after 24 hours transferred to 85% ethanol for 270 YIN ET AL.— NEW PHRUROLITHUS FROM CHINA 271 Figures 1-7. — Phrurolithus bifidus, new species. 1-5. Male. 1. Body, dorsal view. 2. Abdomen, ventral view. 3-5. Palpus. 3. Prolateral view. 4. Retrolateral, sub ventral view. 5. Retrolateral, subdorsal view. 6- 7. Female. 6. Epigynum. 7. Vulva. Scale lines; 1-5 = 1.00mm; 6, 7 = 0.10mm. preservation. Epigyna were cleared in lactic acid for examination and stored in microvials with the specimen. Examination was with an Olympus Tokyo BH=2 stereo dissecting mi- croscope. Leg and palpus lengths are given as: total length (femur, patella + tibia, metatarsus, tarsus). All measurements are in mm. Abbreviations: AER = anterior eye row; AL = abdomen length, ALE = anterior lateral eye, AME = anterior median eye, AME — ALE = distance between AME and ALE, AME — AME = distance between AMEs, AW ~ abdomen width, CH = clypeus height, CL ~ carapace length, CW = carapace width, MOQ = median ocular quadrangle, MOQA — MOQ anterior width, MOQL — length of MOQ, MOQP = MOQ posterior width, PER — posterior eye row, PLE = posterior lateral eye, PME = posterior median eye, PME — PLE = distance between PME and PLE, PME — PME = distance between PMEs, RTA = retrolateral tibial apophysis, TL = total length. Phrurolithus bifidus new species Figs. 1-7 Type material. — Male holotype from sift- ed leaf litter in native forest on pass over Gao- ligongshan, Nankang, 36 air km SE Teng- Chong (24°50'N, 98°47'E, 2100m), Baoshan Prefecture, Yunnan Province, China, 4—7 No- vember 1998, C. Griswold and D. Kavanaugh (deposited in Hunan Normal University, type number 98-NK-48). Paratypes: China: Yun- nan Province: 1 male and 1 female collected with the holotype (female deposited at the Hu- nan Normal University, male at the California Academy of Sciences). Etymology. — The specific name refers to the bifid tibial apophysis of the male palpus. 272 THE JOURNAL OF ARACHNOLOGY Diagnosis. — The male of this species dif- fers from most other east Asian Phrurolithus, except P. vulpes Kamura 2001 and P. pen- natus Yaginuma 1967, in having a bifid tibial apophysis on the palpus. From P. vulpes (see Kamura 2001, figs. 1-5) and P. pennatus (see Paik 1991, figs, 51-54 and Danilov 1999, fig. 3E-F) it differs in having the retrolateral prong of the RTA almost as long as the dorsal prong (as opposed to less than one third the length of the dorsal prong in the other spe- cies), and the dorsal prong apically spatulate (with subapical notch in P. vulpes and apically attenuated in P. pennatus). The female differs from most Asian Phrurolithus in having very large spermathecae and appears most similar to P. hamdeokensis Seo 1988 (see Paik 1991, figs. 29-38) as both species have an epigynum with a single central atrium bearing the cop- ulatory openings and spermathecae which are asymmetrical. It differs from P. hamdeokensis in having a larger epigynal atrium and larger spermathecae that overlap along their anterior margins. Description. — Male.- Carapace pear- shaped, brown with pale gray pattern. AER and PER straight in dorsal view. Cervical groove distinct, head region slightly elevated. Fovea short, longitudinal, anteriorly with bi- lobed pale mark, laterally with radiating mark- ings along radial grooves. Sternum yellow brown, margins darker with gray radial striae inwards. Chelicera weak, grayish brown, with I retromarginal and 3 promarginal teeth. En- dites and labium grayish brown. Palpus and leg segments yellow-brown, distal end of leg femur having grayish black annulus, tibia and metatarsus with one dark stria. Anterior leg spines: 4 on prolateral surface of femur, 7 pairs ventrally on tibiae, and 4 pairs ventrally on metatarsi. Abdomen with dorsum grayish black patterned with yellowish gray, anteriorly with pale cardiac mark and lateral spots, pos- teriorly with five chevrons (Fig. 1); venter yel- low gray with dark markings (Fig. 2). Palpus with large, spherical genital bulb bearing short, curved embolus apically; tibial apophysis large, deeply bifid, v/ith spatulate dorsal and pointed retrolateral prong; femur with small ventral knob about 2/3 the distance from its basal margin (Figs. 3-5). Female.” Coloration and markings as in male, except carapace reddish brown; eye ar- rangement as male. Epigynum with small sub- Figures 8-10. — Phrurolithus qiqiensis, new spe- cies, female. 8. Body, dorsal view. 9. Epigynum. 10. Vulva. Scale line: 8 = 1.00mm; 9, 10 = 0.10mm. circular atrium (Fig. 6); vulva with two large spermathecae from which thin copulatory ducts extend along semicircular arc to atri- um.(Figs. 6-7) Measurements: Male (holotype).” TL 3.90, CL 1.70, CW 1.50; AL 2.10, AW 1.30. Eye sizes and interdistances: AME 0.10, ALE 0.10, PME 0.09, PLE 0.13, AME- AME 0.05, AME- ALE 0.025, PME-PME 0.40, PME-PLE 0.03; MOQL 0.25, MOQA 0.24, MOQP 0.27; CH 0.14, longer than AME diameter. Append- age lengths: palpus: 2.10 (0.50, 0.60, 0, 1.00); leg I: 5.94 (1.68, 2.50, 1.06, 0.70); leg II: 5.50 (1.50, 2.00, 1.30, 0.70); leg III: 4.30 (1.40, 1.50, 0.90, 0.50); leg IV: 6.62 (1.92, 2.00, 1.80, 0.90); leg formula: IV, I, II, III. Female.” TL 4.10, CL 1.60, CW 1.40; AL 2.40, AW 1.40. Appendage lengths: palpus: 1.42 (0.32, 0.60, 0, 0.50); leg I: 4.96 (1.40, 1.71, 1.25, 0.60); leg II: 4.85 (1.35, 1.80, 1.00, 0.70); leg III: 3.70 (1.10, 1.10, 1.00, 0.50); leg IV: 6.10 (1.65, 1.90, 1.70, 0.85); leg formula: IV, I, II, III. Distribution. — Yunnan Province, China. Phrurolithus qiqiensis new species Figs. 8-10 Type material. — Female holotype collect- ed in Qiqi He, 9.9 air km W of Gongshan (27°43' N, 98°34' E, 2000m), Mt. Gaoligong, Nujiang State Nature Reserve, Nujiang Pre- YIN ET AL.— NEW PHRUROLITHUS FROM CHINA 273 fecture, Yunnan Province, China, 9-14 July 2000, HM. Yan, D. Kavanaugh, C.E. Gris- wold, H.-B. Liang, D. Ubick, and D.-Z. Dong (deposited in Hunan Normal University, type number OO-QF-44). Paratypes: China: Yunnan Province: 3 females collected with holotype; Baoshan Prefecture: 1 female (No. 98-OP- 15), pass over Gaoligongshan, Luoshuidong, 28 air km E TengChong, 24°57'N, 98°45'E, 2300m, flight trap in clearing of native forest, 26-31 October 1998, C. Griswold, D. Kavanaugh, and C-L. Long (2 paratypes deposited at the California Academy of Sciences, the rest at Hunan Normal University). Etymology. — The specific name is an ad- jective derived from the type locality. Diagnosis. — This species is similar to Phrurolithus taiwanicus Hayashi & Yoshida 1993 (see Kamura 2001, figs. 14-19) in hav- ing large spermathecae with separate copula- tory openings and large, curved fertilization ducts, but differs in having larger spermathe- cae which reach the epigastric furrow. Description.' — ^Female." (Fig. 8): Carapace pear-shaped, brown-black. AER slightly re- curved, PER straight in dorsal view. Cervical groove distinct and deep, head region some- what elevated. Fovea short, longitudinal, sur- rounded by radiating pale pattern. Sternum heart-shaped, grayish black, paler in center, margins red brown. Chelicera weak, yellow- brown with grayish black mark dorsally, teeth small, 3 promarginal and 2 retromarginal. Pal- pus grayish black interrupted by grayish brown. Eedites and labium grayish black, dis- tal end pale yellow, labium longer than wide. Leg grayish brown, with paired ventral spines. Leg I, tibia with 8, metatarsus with 4, Leg II, tibia with 7, metatarsus with 3. Abdomen dor- sum slightly paler than carapace, pattern yel- lowish gray, cardiac mark not distinct; ante- rior half with 3 pairs markings and 3 chevrons posteriorly; venter with gray markings: 1 pair of longitudinal striae and 2 pairs of patches. Spinnerets pale brown. Epigyeum with large ovoid atrium in pos- terior portion; vulva with two large sperma- thecae and twisted copulatory ducts with thicker anterior and thinner posterior sections (Figs. 9, 10). Male," Unknown. Measurements: Female (holotype).* TL 4.90, CL 1.80, CW 1.50; AL 3.00, AW 2.40. Eye sizes and interdistances: AME = ALE = PME 0.09, PEE 0.10; AME-AME 0.05, AME-ALE 0.03; PME-PME 0. 10, PME-PLE 0.07. MOQ L 0.29 MOQA W 0.20, MOQP W 0.30; CH 0.15, longer than AME diameter. Appendage lengths: palpus: 2.23 (0.78, 1.05, 0, 0.40); leg I: 6.46 (1.70, 2.50, 1.51, 0.75); leg II: 5.35 (1.40, 2.00, 1.20, 0.75); leg III: 4.41 (1.20, 1.51, LOO, 0.70); leg IV: 6.60 (1.70, 2.20, 1.80, 0.90); leg formula: IV, I, II, III. Distribution. — Yunnan Province, China. Phrurolithus revolutus new species Figs. 11-16 Type material. — Female holotype collect- ed at 9 km ESE of Pianma (25°36' N, 98°24' E), Mt. Gaoligong, Yunnan Province, China, 13-18 October 1998, C. Griswold, D. Kavan- augh, and C-L. Long (deposited in Hunan Normal University, type number 98-EP-15). Etymology. — The specific name refers to the curved copulatory ducts. Diagnosis. — This species is similar to Phrurolithus vulpes (see Kamura 2001, figs. 1-7) in having large spermathecae and paired, crescent shaped copulatory openings but dif- fers in having: 1) larger spermathecae which reach the epigastric furrow and 2) the copu- latory openings more posteriorly placed. Description. — Female.- Carapace pale black brown, patterned with black brown. AER and PER recurved in dorsal view. Cer- vical groove distinct, head region slightly el- evated. Carapace with distinct radial grooves, interspersed with shorter lines; fovea short, longitudinal. Sternum heart-like, grayish brown, paler in center, margins red brown. Chelicera weak, yellow-brown, with grayish brown mark dorsally, with 3 promarginal and 2 retromarginal teeth (Fig. 12,). Palpi, endites, and legs yellow-brown, femur with one gray- ish brown band on distal end. Tibiae I, II dark in color, with 6 pairs of ventral spines; meta- tarsi I, II with 4 pairs of ventral spines (Fig. 13); anterior femora with 3 prolateral spines (Fig. 14). Abdomen dorsum iridescent with grayish black and yellow and gray patterns; cardiac mark large, pale grayish black, lined with yel- low-gray margins; sides with fine longitudinal lines; posterior with 2-3 transverse bands and 1 small ellipsoid, white marking; venter me- dian grayish black, laterally with yellow gray oblique striae; spinnerets short, yellow brown. 274 THE JOURNAL OF ARACHNOLOGY Figures 11 — 16. — Phnirolithus revolutus, new species, female. 11. Body, dorsal view. 12. Chelicera, retrolateral view. 13. Leg I prolateral view, showing paired ventral spines. 4. Femur II, prolateral view showing spines. 15. Epigynum. 16. Vulva. Scale bars; 11, 13, 14 = l.OOmm; 12, 15, 16 = 0.10mm. Epigynum with atrium consisting of two small round pits near posterior margin; vulva with 2 large spermathecae and spiraled cop- ulatory ducts (Figs. 15, 16). Male; Unknown. Measurements: Female (holotype); TL 3.35, CL 1.45, CW 1.05; AL 1.90, AW 1.15. Eye sizes and interdistances: AME 0.07, ALE = PLE 0.08, PME 0.55, AME-AME 0.025, AME-ALE 0.028; PME-PME 0.075, PME- PLE 0.05; MOQL 0.20, MOQA 0.14, MOQP 0.18; CH 0.075, almost equal to AME diam- eter. Appendage lengths: palpus: 1.50 (0.60, 0.45, 0, 0.45); leg I: 5.30 (1.35, 2.00, 1.45, 0.50); leg II: 4.42 (1.12, 1.50, 1.10, 0.70); leg III: 3.72 (1.12, 1.20, 0.90, 0.50); leg IV: 5.15 (1.50, 1.60, 1.30, 0.75); leg formula: I, IV, II, III. Distribution. — Yunnan Province, China. ACKNOWLEDGMENTS Thanks go to Prof. Li Heng and Dr. Long Chun-Lin for support for the 1998 and 2000 Sino-American expedition to the Gaoligong Mountains, Drs. Charles Griswold, Jan Bos- selaers, Robert Raven and Mark Harvey for reviewing this manuscript, and Mr. Larrie Currie and the CAS library staff for providing obscure references and citations. This research work was sponsored by the California Acad- emy of Sciences Center for Biodiversity Re- search and Information, the China Natural History Project, and the Foundation of Nature Sciences of Education Department of Hunan Province. LITERATURE CITED Danilov, S.N. 1999. The spider family Liocranidae in Siberia and Far East (Aranei). Arthropoda Se- lecta 7:313-317. Deeleman-Reinhold, C.L. 2001. Forest spiders of South East Asia: with a revision of the sac and ground spiders (Araneae: Clubionidae, Corinni- dae, Liocranidae, Gnaphosidae, Prodidomidae and Trochanterriidae [sic]). Brill, Leiden, 591 pp. Dondale, C.D. & J.H. Redner. 1982. The insects and arachnids of Canada, Part 9. The sac spiders of Canada and Alaska, Araneae: Clubionidae and Anyphaenidae. Research Branch, Agriculture Canada, Publ. 1724:1-194. Hayashi, T & H. Yoshida. 1993. Three new species of the family Clubionidae (Arachnida: Araneae) from Taiwan. Acta Arachnologica 42:47-53. YIN ET AL.— NEW PHRUROLITHUS FROM CHINA 275 Kamura, T. 2001. Seven species of the families Lio- cranidae and Corinnidae (Araneae) from Japan and Taiwan. Acta Arachnologica 50:49-61. Koch, C.L. 1835. Arachniden. In Herrich-Schaffer, G. A. W., Deutschlands Insekten. Heft 128-133. Paik, K.Y. 1991. Korean spiders of the genus Phru- rolithus (Araneae: Clubionidae). Korean Arach- eology 6:171-196. Platnick, N.L 2002. The world spider catalog. Ver- sion 3.0 (updated 24 July 2002). http://research. amnh.org/entomoiogy/ Seo, B.K. 1988. Classification of genus PhrurolP thus (Araneae: Clubionidae) from Korea. Jour- nal, Natural Science Research Institute Seoul 7: 79-90. Song, D.X., M.S. Zhu & J. Chen. 1999. The Spiders of China. Hebei Science and Technology Pub- lishing House, Shijiazhuang, 640 pp. + 4 plates. Song, D.X, & S.X. Zheng. 1992. A new species of the family Liocranidae (Araneae) of China. Si- nozoologia 9:103-105. Wunderlich, J. 1992. Die Spinnen-Fauna der Mak- aronesischen Inseln: Taxonomic, Okologie, Bio- geographie und Evolution. Beitrage fiir Araneo- logie 1:1-619. Wunderlich, J. 1995. Beschreibung der neuen Gat- tung Phrurolinillus der Familie Corinnidae aus Europa (Arachnida: Araneae). Beitrage fur Ara- neologie 4:739-742. Yaginuma, T. 1967. Revision and new addition to fauna of Japanese spiders, with descriptions of seven new species. Literary Department review, Otemon Gakuin University, Osaka 1:87-107. Yin, C.M., X.J. Peng, L.S. Gong & J.P. Kim. 1997. Three new species of the genus Phrurolithus (Araneae: Liocranidae) from China. Korean Ar- achnology 13:25-30. Manuscript received 13 December 2002, revised 6 May 2003. 2004. The Journal of Arachnology 32:276-283 THE MOVEMENT AND ACTIVITY PATTERNS OF SIMILAR-SIZED ADULT AND JUVENILE CRAB SPIDERS MISUMENA VATIA (ARANEAE, THOMISIDAE) Heather L. Sullivan and Douglass H. Morse Department of Ecology and Evolutionary Biology, Box G-W, Brown University, Providence, Rhode Island 02912 USA ABSTRACT. Adult male animals are commonly believed to exhibit higher activity than other conspe- citics, but little information exists to compare their activity with that of other conspecifics of similar size. Here we compare the activity of adult male and similar-sized juvenile female crab spiders Misiimena vatia (Araneae, Thomisidae). Adult males moved farther and more frequently than juvenile females of similar size (fourth instar) that were not affected by impending molt. Juvenile females influenced by impending or recent molt did not move as far or as frequently as nonmolting juveniles, even though their exoskeletons were hard enough to permit rapid movement. A small sample of penultimate males, of similar size to the adult males and juvenile females, exhibited activity patterns similar to the juvenile females. All of these data indicate that the high activity level of adult males is not a simple manifestation of behavior that is solely a function of size. We suggest that the high activity levels of the adult males facilitate search for scarce, cryptic mates. Keywords: Activity level, molt, sit-and-wait predator The high level of activity commonly attri- buted to reproductively active adult male an- imals is typically associated with searching for fertile females, a trait likely to enhance reproductive success by increasing contact with females (Thornhill & Alcock 1983; An- dersson 1994). Seldom, however, is it explic- itly established whether these perceived high activity levels of the males are unique to them. Hypothetically, the size differences of ju- venile and adult animals could by themselves account for their respective levels of activity. Adults of many species considerably exceed the size of younger individuals. However, one way to control the effect of size on activity is to compare the activity of different categories of a species that reach similar sizes (in this instance, adult males, juvenile females, and penultimate males). Size relationships such as the one considered here occur in species char- acterized by small adult males and large, rel- atively immobile adult females. In these sys- tems, scramble competition for finding virgin females (Ghiselin 1974; Parker 2000) may re- sult in high levels of activity by males, poten- tially driven by sexual selection. ' Corresponding author. The crab spider Misumena vatia (Clerck 1757) (Thomisidae) is an excellent species for addressing how activity rates vary over the life cycle. It is a sit-and-wait predator that hunts on flowers for visiting insect prey (Morse 1979; Morse & Fritz 1982). Males are tiny in relation to adult females, at times no more than 1 % of the mass of gravid adult fe- males (Gabritschevsky 1927; LeGrand & Morse 2000). Thus, they provide an opportu- nity to compare the activity of adult males with juvenile females of similar size. Al- though sit-and-wait predators, adult males have shorter giving-up times on hunting sites than adult females, penultimate females (both larger), or penultimate males (similar size) (Morse & Fritz 1982; Chien & Morse 1998; LeGrand & Morse 2000). The difference in activity between adult fe- males and adult males could be merely a con- sequence of the large adult females becoming less mobile than the earlier instars, or adult males becoming more active or both. Given the likely pressures of scramble competition, with adult male movement focused on finding virgin females (Vollrath & Parker 1992; Ko- tiaho et al. 1998), we predicted that adult 276 SULLIVAN & MORSE— MOVEMENT OF CRAB SPIDERS 277 males would move more rapidly (number of moves and distance per unit time) than other life-cycle stages. Since they often do not de- tect sexually mature females at distances ex- ceeding 5 cm (Holdsworth & Morse 2000), male M. vatia should experience especially strong selection to move rapidly, thereby max- imizing their number of visits to sites poten- tially occupied by these females (LeGrand & Morse 2000; Anderson & Morse 2001). Here we make the more critical comparison by test- ing the hypothesis that adult males are more active than other similar-sized conspecifics; i. e., fourth-instar females. We also test wheth- er any such differences involve frequency of moves or length of moves. To obtain the ap- propriate data, we developed a practical meth- od for quantitatively describing movement and activity levels of these spiders. Like all arthropods, spiders spend most of their lives encased in jointed, chitinous exo- skeletons, structures that provide protection and prevent desiccation. However, aside from their soft abdomen, spiders cannot increase in size except by molting, a state that profoundly affects activity and movement (Foelix 1996). Since M. vatia molt frequently (Gabritschev- sky 1927), it was necessary to establish un- equivocally whether the juveniles under study were influenced by impending molt, which could only be accomplished after establishing when an individual molted. It was thus also necessary to test the juveniles at both inter- molt and molt periods, and we consequently present these results and comment upon them as well. Voucher specimens of Misumena vatia have been deposited in the American Museum of Natural History. METHODS Study site and study organisms. — We conducted this study at the Darling Marine Center in South Bristol, Lincoln County, Maine. We performed all trials during June- August of 1998 and 1999 in a 3.5 ha field that is mown yearly in October. Misumena vatia occur naturally at the study site and in surrounding fields and roadsides, spending most of their time in the herbaceous vegetation above 20 cm. They typically course through the fields, moving through most of the vegetation relatively quickly, but remaining on flowers, which they use for hunting sites. for considerably longer periods (Morse & Fritz 1982; LeGrand & Morse 2000). We used juvenile females comparable in size to the adult males. Other than being qualitatively sized by eye, they were randomly collected. We collected the M. vatia used in this study along roadsides adjacent to the study area. We recorded mass, carapace width, and length of leg 1 of each individual at capture. We also noted when a juvenile spider molted and took its new measurements. Except during mea- surements and behavioral trials, spiders were maintained individually in 7 dram clear plastic vials (5 cm tall, 3 cm diameter) and fed mos- quitoes, flies and small moths every third day. Movement and activity. — We observed both free-ranging spiders in the field and also monitored spiders in activity cages set in the field. Cages were 30 X 30 X 30 cm and cov- ered on all sides, including the bottom, with dark green polyester mesh (8 X 10 mesh fi- bers/cm^). We ran all trials in the field in clear weather during daytime hours, the time at which the majority of M. vatia' ^ activity oc- curs (Morse 1979; Morse & Fritz 1982). In initial free-ranging trials we placed a fo- cal spider on a stem of grass approximately 20 cm above the ground. To avoid sites that would slow spider movement, we ran all trials where no flowers were present, thus allowing us to record maximum searching capability, rather than movement confounded with ex- tended visits to flowers. This measure should indicate the maximum distance the spiders would move over a unit of time and should also reflect their ability to locate a favored hunting site or object, be it a flower or a fe- male. Following a 5 min acclimation period, free-range trials lasted 1 h, with activity and location recorded every 5 min, for a total of 12 observation times per individual. Depend- ing upon the spider’s position and activity at the instant of recording, five categories of ac- tivity were recognized: A), location changed since previous observation, in motion at mo- ment observed (either traveling or changing orientation); B). location unchanged, but changing orientation at moment observed; C). location changed since last observation, but stationary; D). location unchanged, but ori- entation changed; E). neither location nor ori- entation changed, stationary. (Categories B, D and E cannot exclude the possibility that a spi- 278 THE JOURNAL OF ARACHNOLOGY der has changed its location but then returned to it.) We had originally intended to invest most of our efforts in measuring free-ranging indi- viduals in the field. However, it quickly be- came apparent that we were losing so many individuals in the open field tests that it was extremely difficult to obtain complete runs of adult males before losing them. In particular, we frequently lost the small agile males in the vegetation and litter in spite of our best efforts to monitor them. Further, individuals lost pre- maturely usually moved more rapidly than those followed for an hour. Given the small sample available, it was not practical to run any of the penultimate males in the open field. The multiple runs needed to ascertain the molt stage of the juvenile females were also im- praetical because of the danger of losing them during one of these runs. We thus gave up the efforts to measure free- ranging individuals in the field and concen- trated on running cage trials. This technique provided a standardized substrate for measur- ing movement, thereby permitting direct com- parison between individuals and between tri- als. The cages allowed us to eliminate hunting sites that would prevent accurate measurement of maximum activity levels. Most importantly, they eliminated the loss of male spiders. We used the same activity categories and times for individuals in the cage trials as for the field trials. We positioned each cage in the open, di- rectly on the ground, and transferred the spi- der from its vial to the middle of the mesh bottom of the cage. Positions of the spiders were determined with a millimeter rule, ori- entation noted, and changes measured and re- corded at each 5 min interval. We thoroughly scrubbed the mesh and rotated the cages be- tween runs to eliminate any possible position effects. Since exhaustive experiments have re- vealed no effect of pheromones on lines or in the air (Anderson & Morse 2001), we did not further control for this factor. We ran a max- imum of four cage trials simultaneously, stag- gering the start by 1 min per cage. Condition of nonmolting and molting in- dividuals.— We divided the runs of the juve- nile females into those affected by molts and those made at intermolt intervals. Molt peri- ods encompassed the time when the integu- ment commenced to break down, leading up to the molt, through the time when the integ- ument of the next instar had completely hard- ened. Premolting changes can often be de- tected as parts of the integument begin to take on a somewhat transparent, vacuolated ap- pearance, the start of the breakdown of the old integument. This change begins 4-5 d before molt (Foelix 1996); we therefore selected, a priori, the period from 5 d before molt to 1 d after molt as the “molting period”, and the rest of the time as the “nonmolting period”. Runs were performed independently of these periods, since we could only make the above- noted separation into nonmolting and molting condition after recording ecdysis. Tests on molting days were run only after the carapace had hardened enough so that the spiders were able to move about readily. All penultimate males had to be tested similarly to the fe- males, and it was the failure of most of these individuals to remain in a nonmolting state over 5 d that resulted in the small sample size. Analysis. — Each adult male and penulti- mate male contributed one activity trial to the analysis. Several juveniles contributed two tri- als, one as a “nonmolter” and one as a “molt- er”. Juveniles were tested daily to determine when they could be incorporated into non- molting and molting categories. Molting in- dividuals were categorized as those between 5 d pre-ecdysis and one day post-ecdysis, fol- lowing Foelix (1996). Only the first nonmolt- ing run and first molting run were used for the analyses. We obtained a measure of total activity for each individual: the number of 5 min periods in which an individual’s activity warranted designation of Category A, B, C or D. A max- imum performance would be 12, a score of 1 being assigned for each of the 5 min obser- vation periods in which movement took place during an hour (Category A, B, C or D). We also calculated a high activity measure (num- ber of Category A movements only) for the cage trials. We reasoned that individuals trav- elling at observation times likely moved more often than those whose positions changed, but were not observed in transit at observation times. We measured movement by summing the distances between an individual’s loca- tions on consecutively occupied sites. This measure gives a minimal possible distance traveled by the spiders; distances traversed during reversals of direction or roundabout SULLIVAN & MORSE— MOVEMENT OF CRAB SPIDERS 279 Table 1. — Total activity scores and distances (± SE) moved by free-ranging spiders. Group n Activity score Distance Adult male 12 8.4 ± 0.58 197.9 ± 62.87 Juvenile female 10 7.0 ± 0.85 1 17.8 ± 25.46 circuits within a single observational period would not be recorded. RESULTS Mass and body dimensions. — Adult males {n ~ 32) weighed 5.2 ± 0.25 mg (jc ± SE) when captured, with carapaces 1 .4 ± 0.03 mm wide and legs 6.3 ±0.13 mm long. Mass was positively correlated with carapace width (r = 0.690, P < 0,001, one-tailed product-moment correlation), and leg length (r = 0.373, P < 0.05, same test). Carapace width and leg length were also positively correlated (r == 0.549, P < 0.01, same test). Sequential Bon- ferroni tests (Rice 1989) were applied to all of the correlations presented in this section. The juveniles {n = 38) weighed 5.2 ± 0.40 mg and measured 1.3 ± 0.04 mm in carapace width and 4.9 ± 0.22 mm in leg length. Mass was positively correlated with carapace width (r == 0.743, P < 0.001, same test) and leg length (r = 0.713, P < 0.001, same test). Car- apace width and leg length were positively correlated (r = 0.843, P < 0.001, same test). Males and females used in these experiments did not differ significantly in either mass {t = 0.090, P > 0.9) or carapace width (r = 0.287, P > 0.5), but did differ in limb length {t = molters non- adult molters males Figure 1 . — Activity levels ± SD for adult males {n = 32), nonmolting juvenile females {n — 38), and molting juvenile females {n = 20). Black bars = total activity, white bars = high activity. 4.324, P < 0.001), all in two-tailed r-tests. Thus, males had longer legs than females of similar size. Free-ranging trials. — We present the data on free-ranging trials to illustrate the difficulty of obtaining velocity measures of adult males in the field and to justify our resort to the cage trials. In the free-ranging trials (Table 1), total activity scores of the adult males did not differ significantly from those of nonmolting juve- nile females {U = 35.5, Z = 1.315, F > 0.05, one-tailed Mann-Whitney f/-test). Neither did the distance traveled differ significantly be- tween the two groups {U = 38, Z = 1.137, P >0.1, same test). However, since both mean activity and mean total distance traveled by the adult males considerably exceeded those of the juvenile females (Table 1), the nonsig- nificant levels were likely a consequence of the especially high variance of the field indi- viduals. Additionally, several rapidly moving males were lost in the field before adequate information could be gathered from them for a measurement. These results prompted our effort to design a method that would eliminate the losses of experimental subjects and that would decrease variance due to likely arti- facts. Cage trials. — Adult males were signifi- cantly more active (total activity scores) than nonmolting juvenile females {U = 394.5, n = 32, 38; Z = 2.517; P < 0.02, one-tailed Mann- Whitney U-test), and the difference in high activity scores between the two groups was particularly large {U = 229.5, Z = 4.462, P < 0.0001, same test) (Fig. 1). Adult males also moved greater distances than nonmolting juvenile females {U = 380, Z = 2.688, P < 0.01, same test) (Fig. 2), a consequence of making both longer and more frequent moves (Figs. 1 & 2). We have not compared the con- tributions of these two variables statistically, because they are unlikely to be independent of each other. Five penultimate males that fit the molting criteria (did not molt for 5 d following a run) 280 Molters Non- Adult molters Males Figure 2. — Total distance moved (minimiim/h) ± SD and mean lengths of single moves in cage trials by adult males, nonmolting juvenile females, and molting Juvenile females. Black bars = total dis- tance, white bars = mean length of single moves. exhibited activity levels similar to those of nonmolting juvenile females (total activity score = 6.3 ± 1.9; high activity score = 2.3 ± 1.0) {U = 19, Z = 0.001, P > 0.9; U = 74.5, Z = 0.001, P > 0.9 in two-tailed Mann- Whitney fZ-tests). We obtained no data on dis- tance moved from these individuals. On the basis of these few individuals it thus does not appear that the adult male — juvenile female differences in activity we have reported are solely a consequence of the sex in question. Activity near the molt. — In the process of establishing the molting state of juvenile fe- males, we obtained considerable information about the activity levels of individuals in molting as well as nonmolting condition. These spiders exhibited a strong relationship between activity level and time before molt (Fig. 3). Both total activity and high activity scores dropped prior to molt, with a rapid re- turn following ecdysis. The total activity score was lowest for individuals tested on their ac- tual day of molt; the high activity score reached its minimum one day prior to molting (Fig. 3). None of the measurements included individuals with a nonfunctionally soft exo- skeleton. Movements of individuals during molt and nonmolt periods differed signifi- cantly in total activity (Fig. 1:7= 20, n = 20, Z = 3.546, P < 0.001, one-tailed Wilcox- on matched pairs signed ranks test), high ac- tivity (Fig. 1: 7 = 32, Z = 2.330, P < 0.02, same test, and total distance traveled (Fig. 2: THE JOURNAL OF ARACHNOLOGY 10 ◄ ^ 4— > =-► prior post Days from molt Figure 3. — Activity levels (total activity in black, high activity in white ± SD) of juveniles across a molting period. Days from molt depicted on hori- zontal axis such that 0 = molt day. Each point rep- resents mean activity (± SD) of trials on molt day or specific number of days before or after molt. 7 = 15, Z = 3.733, P < 0.001, same test). This difference is a consequence of both the length and number of moves made by these individuals (Figs. 1 & 2). The mean intermolt period was 20.6 ± 6.6 d (Fig. 4). Given a molt phase of approxi- mately 6 d, a juvenile crab spider spent about one-third of its juvenile life in a molt phase, and about one-third of the juveniles were thus in a molt phase at any given time. The length of an instar did not correlate significantly with size (carapace width) (7 = 47, n — 14, Z = 0.345, P > 0.7, two-tailed Wilcoxon matched pairs signed ranks test), nor did instar length 5 10 15 20 25 30 Instar length (d) Figure 4. — Frequency distribution of time in days (d) between molts of juveniles. SULLIVAN & MORSE— MOVEMENT OF CRAB SPIDERS 281 correlate with the percentage of increase in size achieved via molt (T = 43, n = 14, Z = 0,534, P > 0,5, same test) or length of the preceding instar (T = 19, « - 8, Z — 0.140, P > 0.8, same test), DISCUSSION Animals with long, slender legs are often cursorial. Therefore, LeGrand & Morse (2000) hypothesized that, given the differenc- es between juvenile females and penultimate males, the long legs of adult males facilitate rapid movement. In fact, the adult males in this study made significantly longer moves than did juvenile females. However, a sub- stantial part of the difference between adult males and the other groups lay in the frequen- cy of their moves, especially high-activity moves, rather than simply in the length of their m.oves. Thus, the difference in adult male movement cannot be attributed solely to dif- ferences in the shape of their legs. Although one might argue that the cages provided an unnatural environment in which to run the experiments, the distances traveled by the spiders in the cages and in the open were similar (though less variable). The cages also exposed all of the spiders to the same conditions and cues. We therefore conclude that the cage trials served as an adequate es- timate of movement rates of the different age and sex categories of these spiders in the field. Innate behavioral factors might dictate the initiation of movements, as seen in the initi- ation of other behavioral patterns (Morse 2000) of M. vatia, but physiological mecha- nisms may dictate the length of the move- ments in part. Since spiders are physiologi- cally severely limited in their ability to generate aerobic activity (Linzen & Gallowitz 1975; Anderson & Prestwich 1982), the adult males may avoid severe oxygen debts by re- stricting the lengths of their movements to the modest ranges observed, even though they ex- ceeded those of the juvenile females. Even highly cursorial wolf spiders (Lycosidae) have tightly constrained sprint distances (Bristowe 1939; Morse 1997). In other spiders investigated, females usu- ally release pheromones prior to mating (e. g., Tietjen & Rovner 1982; Fernandez-Montrav- eta & Ruano-Bellido 2000), behavior that at- tracts males, as a result of following lines ''scented” with pheromones or, possibly, by airborne pheromones (Searcy et al. 1999). However, adult male M. vatia only detect fe- males within a limited range of a few cm (LeGrand & Morse 2000) and follow drag- lines somewhat indiscriminately (Anderson & Morse 2001). Although the females are ca- pable of mating immediately following molt (Holdsworth & Morse 2000), adult females in our populations on average were not mated until two to three days after molting, probably because, in the apparent absence of cues, the males could not locate them quickly (LeGrand & Morse 2000). The two-three day hiatus be- tween molt and mating suggests that sizeable reproductive opportunities may be available to agile individuals. Therefore, males should be under extremely strong selective pressure to move frequently, quickly, and efficiently in order to find females. The high activity levels of the males should facilitate search for these cryptic females. We initially planned extensive compari- sons between adult males and penultimate males, which are of similar size, but penul- timate males were available in the field for only limited periods in both the autumn and spring. We did not run them in the autumn out of concern that impending diapause might bias the results (Tauber et al. 1986; Ta- naka 1992). In the spring most penultimate males molted within five days of their initial test and thus could not be used for this com- parison, since before certifying a run for use in the analysis we had to establish that they were in a nonmolting state. The results from the small sample agree with the more general earlier findings of activity among penultimate males (LeGrand & Morse 2000) and strengthen the conclusion that the unique rates of adult male movement are not a mere consequence of sex, but are related directly to male maturation. Juveniles 5 d prior to molt through 1 d after molt were significantly less active than non- molting juveniles of the same size, whether using activity scores or distance traveled as criteria. Spiders tend to withdraw from sight and refuse prey for up to nearly a week prior to ecdysis (Foelix 1996). The initial decrease in activity of molting juveniles was probably a consequence of the onset of apolysis (sep- aration of the new epidermis from the old cu- ticle). The commencement of apolysis is dif- ficult to predict, but may precede ecdysis by 282 THE JOURNAL OF ARACHNOLOGY up to a week in insects and spiders (Wiggles- worth 1984; Foelix 1996). Differences in ac- tivity between molting and nonmolting juve- niles resulted both from the molters moving less frequently and exhibiting far less high- activity behavior than nonmolting individuals. In contrast, the greatest difference between adult males and both molting and nonmolting females lay in the large amount of high-activ- ity behavior in the adult males. Molting-con- dition juveniles thus exhibit severely con- strained activity levels, which result in large part from a decline in the number of moves they make, rather than the length of moves. It must be emphasized that this low level of ac- tivity is not a consequence of a physical in- ability to move, since the period of forced in- activity resulting from a soft exoskeleton constitutes only several hours of the six-day period of low activity. ACKNOWLEDGMENTS We thank J. Anderson, S. Chien, R. Eeld- man, A. Holdsworth, H. Hu, J. Kraus, A. Ko- pelman, and R. LeGrand for assistance and J. K, Waage for comments on the manuscript. We also thank K. J. Eckelbarger, T. E. Miller, and other staff members of the Darling Marine Center of the University of Maine for facili- tating fieldwork on their premises. In partic- ular, T.E. Miller helped in many ways to ex- pedite research at the site. HLS's work was supported by a Howard Hughes Undergradu- ate Summer Eellowship. Partially supported by the National Science Eoundation IBN98- 16692. LITERATURE CITED Anderson, J.E & K.N. Prestwich. 1982. Respiratory gas exchange in spiders. Physiological Zoology 55:72-90. Anderson, J.T. & D.H. Morse. 2001. Pick-up lines: how male crab spiders find reproductive females. Behavioral Ecology 12:360-366. Andersson, M. 1994. Sexual Selection. Princeton University Press, Princeton, New Jersey. Bristowe, W.S. 1939. The Comity of spiders. Ray Society, London. Chien, S.A. & D.H. Morse. 1998. Foraging patterns of male crab spiders Misumena vatia (Araneae, Thomisidae). Journal of Arachnology 26:238- 243. Fernandez-Montraveta, C. & J. Ruano-Bellido. 2000. Female silk and mate attraction in a bur- rowing wolf-spider (Araneae, Lycosidae). Bul- letin of the British Arachnological Society 1 1 : 361-366. Foelix, R.E 1996. Biology of Spiders, 2"^ ed. Ox- ford University Press, New York. Gabritschevsky, E. 1927. Experiments in color change and regeneration in the crab spider, Mis- umena vatia. Journal of Experimental Zoology 47:251-267. Ghiselin, M.T 1974. The Economy of Nature and the Evolution of Sex. University of California Press, Berkeley, California. Holdsworth, A.R. & D.H. Morse. 2000. Frequen- cies of mate guarding and female aggression in the crab spider Misumena vatia. American Mid- land Naturalist 143:201-211. Kotiaho, J., R.V. Alotalo, J. Mappes, S. Parri & A. Rivero. 1998. Male mating success and risk of predation in a wolf spider: a balance between sexual and natural selection? Journal of Animal Ecology 67:287-291. LeGrand, R.S. & D.H. Morse. 2000. Factors driving extreme sexual dimorphism of a sit-and-wait predator under low density. Biological Journal of the Linnean Society 71:643-664. Linzen, B. & R Gallowitz. 1975. Enzyme activity patterns in muscles of the lycosid spider Cupien- niiis salei. Journal of Comparative Physiology 96:101-109. Morse, D.H. 1979. Prey capture by the crab spider Misumena vatia (Araneae: Thomisidae). OecoL ogia 39:309-319. Morse, D.H. 1997. Distribution, movement, and ac- tivity patterns of an intertidal wolf spider Par- dosa lapidicina population (Araneae, Thomisi- dae). Journal of Arachnology 25:1-10. Morse, D.H. 2000. Flower choice by naive young crab spiders and the effect of subsequent expe- rience. Animal Behaviour 59:943-951. Morse, D.H. & R.S. Fritz. 1982. Experimental and observational studies of patch-choice at different scales by the crab spider Misumena vatia. Ecol- ogy 63:172-182. Parker, G.A. 2000. Scramble in behaviour and ecol- ogy. Philosophical Transations B, 355:1637- 1645. Rice, W.R. 1989. Analyzing tables of statistical tests. Evolution 43:223-225. Searcy, L.E., A.L. Rypstra & M.H. Persons. 1999. Airborne chemical communication in the wolf spider Pardosa milvina. Journal of Chemical Ecology 25:2527-2533. Tanaka, K. 1992. Photoperiodic control of diapause and climatic adaptation of the house spider, Achaearanea tepidariorum (Araneae, Theridi- idae). Functional Ecology 6:545-552. Tauber, M.J., C.A. Tauber & S. Masaki. 1986. Sea- sonal Adaptations of Insects. Oxford University Press, New York. Thornhill, R. & J. Alcock. 1983. The Evolution of SULLIVAN & MORSE— MOVEMENT OF CRAB SPIDERS 283 Insect Mating Systems. Harvard University Press, Cambridge, Massachusetts, Tietjee, WJ. & J.S. Rovner. 1982, Chemical com- munication in lycosids and other spiders, Pp, 249-279. In Spider communication (P.N. Witt & J.S. Rovner, eds.). Princeton University Press, Princeton, New Jersey. Vollrath, F. & G.A. Parker. 1992. Sexual dimor- phism and distorted sex ratios in spiders. Nature 360:156-159. Wigglesworth, VB. 1984. Insect physiology, 8* ed. Chapman & Hall, London. Manuscript received 3 February 2003, revised 8 September 2003. 2004. The Journal of Arachnology 32:284-312 THE SYSTEMATICS OF THE EREMOBATES SCABER SPECIES-GROUP (SOLIFUGAE, EREMOBATIDAE) Jack O. Brookhart and Paula E. Cushing: Denver Museum of Nature & Science, 2001 Colorado Blvd., Denver, Colorado 80205-5798 USA. E-mail: joipbroo@ix.netcom.com ABSTRACT. The scaber group of the genus Eremobates is reviewed in terms of new characters and a more restricted biogeographic area. Three new species are described from the U.S.A.: Eremobates socal (California), E. icenoglei (California), E. corpink (Utah). We synonymize E. gladiolus Muma with E. scaber (Kraepelin); E. consors Muma, E. ascopulatus Muma and E. flavus with E. ascopulatiis Muma; and E. mimbrenus Muma with E. monuoniis (Roewer). Eremobates scaber, E. hodai Muma, E. clams Muma, E. similis Muma are now described from both sexes. All scaber species except the Mexican species, E. legalis Harvey, are now known from both sexes. We also present the first phytogeny of the species group based on morphological characters. This phytogeny demonstrates a geographic grouping into north- ern and southern clades. Keywords: Solifugids, phytogeny, cladistic analysis, biogeography Muma (1951) redescribed the genus Ere- mobates Banks 1900 to include those solifu- gid species characterized by males with a ba- sally dilated mesoventral groove running the length of the mesodorsal or mesoventral sur- face of the fixed finger. The male flagellum complex consists of dorsal striate setae, ven- tral striate setae, and a flattened apical plu- mose seta covering much of the mesoventral groove. The female genital opercula varied depending upon the species-group. Addition- ally the Eremobates scaber species group was erected by Muma (1951) to include those Er- emobates whose males were characterized by a broad basal notch occupying one-third or more of the length of the fixed finger in dorsal view (Fig. 1). In the scaber group the meso- ventral groove is deep and narrow. The female genital opercula are roughly triangular with species distinguished by differences in the medial margins. Muma (1951) listed six species in the sca- ber group including the typical or defining species E. scaber (Kraepelin 1899) although he had not seen the female type specimen. Af- ter examining type specimens in both the U.S. and Europe as well as other specimens from various collections and collectors Muma (1970) recalled Kraepelin’s 1899 description of Datames seedier based on the female type from “Washington Territory” as the correct characterization of Eremobates scaber. He used his 1951 description, erroneously attri- buted to E. scaber, to establish E. septentrion- is Muma 1970, used his 1951 description of E. geniculatus to erect E. mormonus (Roewer 1934) and defined E. geniculatus (C.L. Koch 1842) (Simon 1879, misidentified) using Si- mon’s 1879 description of a single female from Mexico (Muma 1970). In 1989 Muma described six new species. This resulted in 15 species in the scaber group with E. scaber, E. actenidia Muma 1988, E. clarus Muma 1989, E. consors Muma 1989, E. ascopulatus Muma 1951 and E. hodai Muma 1989 described from only one sex although Muma included the male of E. scaber in the key. Eremobates clarus, E. actenidia and E. consors were each described from a single specimen and E. as- copulatus from two males. Eremobates similis (Muma 1951) was noted as being described in both sexes (Muma 1989), but the female de- scription has not been found. In describing E. scaber, Muma (1951) used specimens from an area that extended from the northwestern United States to Las Vegas, Nevada but noted that it might include other species of this group. In addition, other spe- cies of this group seemed to have sympatric ranges (Muma 1951, 1962, 1989). In each of 284 BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 285 Figures 1-3. — Diagnostics used in compiling data for scaber group. 1 . Eremobates scaber, dorsal view of male fixed finger (arrow identifies basal notch). 2. Diagram of male chelicera showing rang- es of measurement and morphological characters: CL = cheliceral length, CW = cheliceral width, FL = fond length, FW = fond width, FFW = fixed finger width, FF = fixed finger, MF = movable fin- ger, PT = primary tooth, IT = intermediate tooth, AT = anterior tooth. 3. Diagram of female genital operculum. his publications Muma (1951, 1962, 1989) cit- ed several problems with the distinction be- tween species and problems of sympatric as- sociations. Muma (pers. comm.) indicated that this group needed to be more thoroughly stud- ied. For the most part, this group is an inhabi- tant of pifion pine-juniper or desert shrub communities. Muma (1963) identified E. zinni (Muma 1951), E. similis, E. ctenidiellus and E. mormonus as inhabitants of the Mercury, Nevada Nuclear Test Site, a Mojave Desert region, although some of the specimens were misidentified. Allred & Muma (1971) listed E. septentrionis and E. ctenidiellus as inhabitants of the Snake River Plain which is part of the Columbian Plateau. Brookhart (1972) found E, mormonus, later changed to E. similis, in the San Luis Valley of Colorado and E. cten- idiellus in the mesa regions of western Colo- rado. The Sevilleta Long Term Ecological Re- serve project at the northern tip of the Chihuahuan Desert surveyed six distinct de- sert grassland/high desert areas and found E. similis in only the pinon-juniper association (Brookhart & Brantely 2000). At the Hanford Nuclear Site, Rich Zack’s E. scaber material (WSU) was collected in Great Basin Desert shrub habitat, and various Canadian speci- mens were collected in the sagebrush of the Okanogon Valley. Eremobates scaber group species have been collected at 2394 m in Wy- oming, 2303 m in the San Luis Valley of Col- orado, and on Mt. Palomar, California. Muma (1951, 1962, 1970, 1989) used length vs. width of the fondal notch, number and shape of ctenidia, and number of palpal papillae, as well as coloration of appendages to separate each species. The number of cte- nidia ranged from 0-6. The palpal scopula varied from none to over 120 papillae. Fe- males were identified by the structure of the genital operculum and the coloration of ap- pendages. Coloration of eye tubercle and mal- leoli were noted but were consistently the same for all species with eye tubercles dark and malleoli white. Abdominal coloration var- ied from a pale yellow to a grey background dorsally and ventrally with lighter pleural membranes between species and also between specimens of the same species. Many speci- mens had tergites with a rectangular, brown- ish, violet pigmentation which gave the ap- pearance of a broad stripe to many specimens. Muma (1951) calls this a sclerite although it is not particularly thick or hardened. It was not found to be diagnostic in this study. Male chelicerae have no teeth on the fixed finger and some variation in the shape of the fixed finger in ectal view. The movable finger follows the general pattern of a large primary tooth, two intermediate teeth, the posterior being larger and an anterior tooth. The mesal tooth varies from tiny to absent. Female che- licerae have a fixed finger with teeth ordered successively posterior to anterior, intermedi- 286 THE JOURNAL OF ARACHNOLOGY ate tooth, large primary tooth, two interme= diate teeth, medial tooth, a single intermedi- ate tooth and a smaller anterior tooth. The female movable finger has a large primary tooth, a variable sized anterior tooth and two intermediate teeth, the posterior of which is larger. The mesal tooth varies from absent to medium size. Fondal teeth in both male and female grade out I, III, II, IV in size, al- though in some species the fondal tooth III is equal in size to fondal tooth I. Due to wear, the intermediate teeth on both male and fe- male movable fingers are sometimes hard to diagnose. METHODS Because of their nocturnal habits solifug- ids are usually not collected in abundance and study specimens are difficult to obtain (Muma 1951, 1970, 1989; Punzo 1998). Ef- forts were made to accumulate at least five males and five females from identifiable geo- graphical regions. The Sevilleta Long Term Ecological Research (LTER) site provided specimens over a seven year period (Brook- hart & Brantley 2000) that enabled us to identify variations within an isolated species population. Brookhart conducted limited pit- fall projects in the San Luis Valley, the north- west corner of Colorado near Dinosaur Na- tional Monument and the area around Colorado National Monument on the Colo- rado Plateau from 1997-1999. He also col- lected from southeast Utah using the same method during 2000-2001. A one year pitfall series from Hanford Site, Benton County, Washington collected by Rich Zack et al, was also examined. Collections from other insti- tutions were also provided for study. We were able to examine all of the types. Coor- dinates for some specimens are approximate as they are based on historical locale infor- mation. We determined collection coordi- nates for specimens post hoc when possible. However, if the specimen was collected 6 km or more from the given site, the coordinates were not reported since the locale informa- tion on the collection label was considered too vague. Using the methods of Muma (1951), Brookhart & Muma (1981, 1987) and Muma & Brookhart (1988) we measured the length of palpus, leg I, leg IV; length and width of chelicera and propeltidium; length and width of fondal notch; and width of base of fixed finger. Fig. 2 indicates measurement areas. Abbreviations used to indicate various che- liceral structures are as follows: FF = fixed finger; MF = movable finger; PT = primary tooth; AT = anterior tooth; MT = medial tooth; IT — intermediate tooth; MST = mesal tooth. No satisfactory measurement of the depth of the notch could be found but depth could be inferred based on the amount of “crimping” or upturn of the male fixed fin- ger. Those with a deeper notch demonstrated a more deeply crimped aspect when viewed ectally (see ectal view of E. scaber. Fig. 31). Female length; length of palpus, Leg I, Leg IV; length and width of chelicera, propelti- dium and genital operculum were also made. All measurements are in millimeters. The number, shape, and relative length of ctenidia to succeeding tergite was noted. Counts were made of palpal papillae. Color of palpus, legs I, II, III, IV and general overall color especially that of the propeltidium was recorded. The shape of the female genital operculum especially the medial margin was observed using some new terminology (see Fig. 3). In addition to previously utilized charac- ters, the shape of the anterior tooth of the male fixed finger, the absence or presence of a cleft anterior to the anterior tooth on both male and female cheliceral movable fingers, the presence and size of the mesal tooth, the position of the posterior intermediate tooth on the principal tooth of the male and female movable finger were analyzed. We investi- gated the Ectal Cheliceral Cluster Setae (ECCS) of Muma (1985) but found no useful parameters. Ratios used previously by Muma (1951, 1970, 1989), Brookhart & Muma (1981, 1987), Muma & Brookhart (1988), and Brookhart & Cushing (2002) were computed. These ratios are as follows: A/CP = the sum of the lengths of palpus, leg I, and leg IV divided by the sum of length of chelicera and propeltidium indicating length of appendages in relation to body size. The larger the num- ber, the longer legged is the species. FL/FW indicates whether the chelicera fondal notch is longer or wider. Longer is defined as the anterior to posterior axis and width is defined as the dorsal to ventral axis. FW/FFW diag- noses the size of fondal notch compared to BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 287 Figures 4-9. — Eremobates icenoglei, new species. 4, Dorsal view male propeltidium. 5. Ectal view male right chelicera. 6. Mesal view male right chelicera. 7. Ventral view male ctenidia. 8. Ectal view female right chelicera. 9. Ventral view female genital operculum. Scale lines = 1 mm. 288 THE JOURNAL OF ARACHNOLOGY the thickness of fixed finger. CW/FFW is used to indicate whether the fixed chelicera finger is thin or robust in relation to the size of the chelicera. GOL/GOW demonstrates the relative size of the female genital oper- culum in terms of length and width. Because of limited and unequal sample size the Tukey-Kramer Analysis of Variance (AN- OVA) was used to test differences in ratio means of each population as recommended by Sokal and Rohlf (1981). New parameters are given whenever possible for both males and females since Muma’s measurements were sometimes based on a few specimens and from widely separated localities suggesting that more than one species might be included. We attempted to use specimens representative of a given species from a defined geographic province since it appeared that there was no sympatric association except in the Nevada Test Site area where E. zinni and E. ciscopu- latus appear to be sympatric. Over 250 specimens were used in this study, many more than were available to other investigators. We describe the male of E. scci- her and the females of E. clams, E. similis, E. acteuidia and E. hodai. Every species ex- cept E. legcdis Harvey 2002 is now known from both sexes and a reduced and more spe- cific geographic range for each implied. The following institutions have loaned us types and specimens for observations: American Museum of Natural History (AMNH), Nomi Platnick; Athabasca University (ABU), Robert Holmberg; Brigham Young University (BYU), Richard Bauman; California Academy of Sci- ence (CAS), Charles Griswold; Colorado State University (CSU), Boris Kondratieff; University of Colorado at Boulder (CU), Virginia Scott; Florida State Collection of Arthropods (FSCA), G.B. Edwai'ds and Paul Skelly; University of New Mexico Museum of Southwestern Biology, Sandy Brantley; Utah State University (USU), Wilford Hansen; State of Idaho Dept, of Envi- ronmental Quality (ID), WH. Clark; Spencer Museum, University of British Columbia, Van- couver (SMUBC); Royal British Columbia Mu- seum, Victoria (RBCM); Royal Ontario Muse- um, Toronto (ROM); Washington State University James Entomological Collection (WSU), Richard Zack; Museum National d’Histoire Naturalle, Paris, France (MNHN), Christine Rollard. SYSTEMATICS Family Eremobatidae Kraeplin Genus Eremobates Banks 1900 Datames Simon 1879:113 (preoccupied). Datames scaber Kraepelin 1 899. Eremobates Banks 1900:426 (new name for Data- mes Simon). Eremoperna Roewer 1934:557 (in part). Eremopiis Roewer 1934:561 (in part). Eremognatha Roewer 1934:566 (in part), Eremocosta Roewer 1934:569 (in part). Eremostata Roewer 1934:571 (in part). Type species. — Gluvia cinerascens E.L. Koch 1842 (junior synonym of Galeodes pal- lipes Say 1923). Muma (1951) described the genus Eremobates as small to medium sized Eremobatidae with a mesoventral groove that extends the entire length of the male fixed fin- ger. The flagellum complex is composed of a dorsal row of simple tubular bristles that are sometimes striate and a ventral row of S- shaped, flattened, plumose bristles that form an arch over the basal third of the mesoventral groove. The apical, plumose bristle of the ven- tral row is straight and forms a parallel cov- ering over the apical two-thirds of the meso- ventral groove. The first post-spiracular abdominal stern ite of males are with or with- out ctenidia. Genital operculum of female var- iable. This description did not change in later works (Muma 1962, 1970, 1989). Eremobates scaber (Kraepelin 1901) Figs. 31, 42, 47, 52 Datames scaber Kraepelin 1899:243, fig. 19f. Eremobates scaber (Kraepelin 1899), Kraepelin 1901:124-125, fig. 91; Muma 1970:12, fig. 10; Muma 1989:8—9. Eremostata scabra (Kraepelin), Roewer 1934:124. Eremobates gladiolus Muma 1951:57-58, figs. 58- 60; Muma 1970:11-12; Muma 1989:9. NEW SYNONYMY. Type. — Female holotype of Datames sca- ber No. 9137, from “Washington Territory”, US in the E. Simon collection, Paris, France (MNHN). Male holotype of Eremobates glad- iolus from Maupin, Wasco County, Oregon, US (45°1UN, 12U04'W), 19 July 1934, J.M. Pearson, in AMNH. Female allotype from Starbuck, Columbia County, Washington, US (46°3UN, 118°07'W), 4 July 1938, C.S. Bren- ner, in AMNH. Diagnosis. — Males.- Most easily identified by the strongly upturned or “crimped” por- BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 289 16 Figures 10-16. — Eremobates socal new species. 10. Dorsal view male propeltidium. 11. Ectal view right male chelicera. 12. Mesal view right male chelicera. 13. Ventral view male right palpus. 14. Ventral view male ctenidia. 15. Ectal view right female chelicera. 16. Ventral view female genital operculum. Scale lines = 1 mm. 290 THE JOURNAL OF ARACHNOLOGY tion of the fixed finger of the male chelicera (Fig. 31). Two short, thin to flat ctenidia. It is separated from E. clarus by the shape of fixed finger and distinctive female genital opercu- lum. Description. — Males.- Chelicera, propelti- dium and appendages dusky yellow with the following markings: propeltidium blotched dusky purple except for a medial lighter dusky ovoid region (Fig. 47). Some specimens have light violet markings on distal third of leg IV and a chelicera with a dorsal and two lateral dusky purple stripes, abdomen dusky grey. Chelicera as Muma (1951, fig. 58). FF se- verely crimped in ectal view (Fig. 31), MF with PT large, AT small, sharp, triangulate, 2 IT, 1st IT separated from PT, cleft under AT, mesal tooth absent, FT graded I, III, II, IV, FT III triangulate and large as FT I, FL/FW equal to or slightly wider, flattened apical plumose bristle occupies 75-80 percent of mesoventral groove, palpal metatarsus with a scopula of 33-80 rounded papillae; 2 short, thin to flat ctenidia (Fig. 42). Male measurements(5): Hanford Site, Ben- ton County, Washington, (WSU). Total length 18.5-25.0, chelicera length 5.00-6.72, chelic- era width 2.64-3.04, propeltidium length 2.83-3.33, propeltium width 3.96-4.38, pal- pus length 15.5-18.0, 1st leg length 13.0- 16.0, 4th leg length 22.0-26.0. Ratios: A/CP 5.17-6.73, CL/CW 1.84-2.40, FL/FW 0.82- 1.07, FW/FFW 1.28-1.55, CW/FFW 4.86- 7.08, PL/PW 0.70-0.80. Females: Coloration same as males, chelic- era typical of species; FF with PT and MT large, a small AT, two IT between PT and MT, one small IT posterior to the AT; MF with large PT, pointed AT; two small IT, the prox- imal of which is larger; posterior IT separate from primary tooth; no cleft under AT of MF, MST indistinct or absent. Genital operculum as in fig. 10, p. 19, (Muma 1970) with short, thin arms, medial margin lobed, gently re- curved wings ending in a curved posterior margin (Fig. 52). One specimen with 57 rounded papillae on metatarsus of palpus, the rest had none; two tiny hairlike ctenidia were present on one specimen, the rest had none. Female allotype: Total length 19.0, chelic- era length 6.4, cheliceral width 3.1, propelti- dium length 2.9, propeltidium width 4.16, pal- pus length 16.0, first leg length 11.0, fourth leg length 22.5. Ratios: A/CP 5.32, CL/CW 19 Figures 17-19. — Eremobates hodai Muma. 17. Ectal view female chelicera. 18. Ventral view fe- male genital operculum. 19. Ventral view right male palpus. Scale lines = 1 mm. 2.06, PL/PW 7.00, GOL/GOW 0.72. Female measurements (3): Length 19.0—23.0, chelic- era length 6.04-7.50, chelicera width 2.29- 3.29, propeltidium length 2.58-3.30, propeL tidium width 3.96-4.58, palpus length 14.0- 17.0, first leg length 11.0-15.0, fourth leg length 18.0—25.0. Ratios: A/CP 4.86-5.26, CL/CW 2.28-2.95. PL/PW 0.65-0.73, GOL/ GOW 0.50-0.67. Remarks. — Muma (1951) described E. scaber using a sample population of both males and females from a geographic area ex- tending from Washington state to the deserts of Arizona. He remarked as to the variability of the species and suggested that it might in- clude two or more species. Unfortunately his drawings of the female genital operculum (1951, p. 53, fig. 53) were obviously of an- other species. After viewing Kraepelin’s fe- BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 291 male type from Washington Territory, Muma (1970) described E. scaber based solely on the type and indicated that males were unknown. In the same publication (Muma 1970) used his 1951 description of E. scaber to establish the new species, E. septentrionis (Muma 1970). Muma’s (1951) description of E. gladiolus listed the male holotype from Maupin, Oregon, the female paratype from Starbucks, Washington, and paratypes from Umatilla, Oregon and Wishrum, Washington, all in the Columbia River Basin. Muma uses only a slight difference in the coloration on leg IV to differentiate the two species. After examina- tion of material from Hanford Test Site in Washington state and Canadian samples from the Okanogan Valley, and the subsequent ex- amination of the types of E. scaber and E. gladiolus, we have synonymized the two spe- cies under E. scaber based on the shape of the female genital operculum and the male fixed finger. The collection sites indicate a range that en- compasses the Columbian River Basin, and the Okanogan Valley in northern Washington, USA and the Okanogan Valley, southern Brit- ish Columbia, Canada which are primarily high desert shrub communities (USEPA 1986). Specimens examined. — Males.- UNITED STATES: Oregon: Washington County, Mau- pin (45°10'N, 121°04'W), July 1934, J. M. Pierson (d, AMNH); Umatilla County, Uma- tilla (45°55^N, 119°20'W), 24 June 1882, S. Henshaw ($, AMNH). Washington: Benton County, Hanford Nuclear Site (46°32'N, 119°3UW), 23 July-8 August 1999, Rich Zack (8 d, 3 9, WSU); Whitman County, Wawsweiko Peak (46°32'N, 118°79'W), 27 July 1981, no collector data (d, WSU). CAN- ADA: British Columbia: Osyoos, Haynes Ecological Reserve (49°07'N, 119°40'W), 19 June 1986, S.G. Cannings (2 d, Spencer Mu- seum, University of British Columbia); 14 June-3 August 1987 (d, 9, ABU); Osyoos, Mount Kobau, 10-33 July 1991, D. Blades & C. Maier (d. Royal British Columbia Muse- um, Victoria); Penticton (49°10'N, 119°3UW), 1973, W. D. Charles (d, ABU); 5 July 1973, M. Redivo (d ABU); 27 August 1972, Jose Matias (d, ABU); Summerland (49°36'N, 119°40'W), 3 July 1928, T. B. Kurta (9, Roy- al Ontario Museum, Toronto); August-No- vember 1982, W. D. Charles (d, ABU); Ker- emeos (49°12'N, 119°50'W), 6 September 1960, Philip Desjardins (9, Spencer Museum, University of British Columbia); Oliver (49°1UN, 119°33'W), J. Slack (9, Spencer Museum, University of British Columbia). Eremobates ctenidiellus Muma 1951 Figs. 32, 38, 48, 55 Eremobates ctenidiellus Muma 1951: figs. 58-60; Muma 1962:3; Muma 1963:1; Muma 1970:10; Muma & Allred 1971:165; Brookhart 1972:33; Muma 1987:20. Type . — Male holotype and female allotype from 3.2 km west of Glenwood, Sevier Coun- ty, Utah, US (38°45'N, lir59'W) 30 June 1940, Gertsch & Hook in AMNH. Muma (1951) lists paratypes in U.S. National Mu- seum, Museum of Comparative Anatomy, USU, Cornell University (Muma 1951) but there are no paratypes at USU (Wilford Han- sen pers. comm.). Diagnosis. — Pale species, with 0-2 very thin hairlike ctenidia. Posterior IT of male and female ME in the notch of PT, no cleft under AT. Female genital operculum distinctive with virtually no modification of the interior edge except for a heavily chitinized region which makes it appear as if there is a spot on the medial edge. Fondal notch length equal to width. Description. — Males.- Muma’s (1951) de- scription is accurate. Appendages pale yellow, propeltidium dusky, violet brown except for a medial pale yellow ovoid region (Fig. 48), ab- domen dusky. Chelicera as in fig. 58, p.56, Muma (1951). A thin FF slightly crimped in ectal view, ME with large PT and smaller slightly crumpled AT, no cleft anterior to AT, IT in notch of PT. MST tiny to absent, fondal teeth typical (Fig. 32). Palpus with 50-100 rounded, white papillae on the metatarsal scopula, ctenidia 0-2 thin, hair-like setae (Fig. 38). Male holotype: Total length 22.0, chelicera length 5.6, chelicera width 3.2, propeltidium length 3.0, propeltidium width 4.0, palpus 18.0, 1st leg 15.0, 4th leg 23.0. Ratios: A/CP 6.36, CL/CW 3.20, PL/PW 0.75, FL/FW 1.01, FW/FFW 1.25, CW/FFW 5.70. Male mea- surements (5): Total length 17.0-23.0, chelic- era length 4.6-6. 1, chelicera width 2. 2-3. 2, propeltidium length 2. 4-3. 2, propeltidium width 3. 3-4.4, palpus length 15.0-18.5, first leg length 13.0-15.0, fourth leg length 17.5- 292 THE JOURNAL OF ARACHNOLOGY Figures 20-26. — Eremobates corpink new species. 20. Dorsal view male propeltidium. 21. Ectal view male right chelicera. 22. Mesal view male right chelicera. 23. Ventral view male right palpus. 24. Ventral view male ctenidia. 25. Ectal view female chelicera. 26. Ventral view female genital operculum. Scale lines = 1 mm. 25.0. Ratios: A/CP 6.15-7.23, CL/CW 1.64- 2.34, PL/PW 0.58-0.80, FL/FW 0.80-1.00, FW/FFW 2.67-3.75, CW/FFW 5.50-7.00. Females: Coloration the same as in the males. Chelicera typical of species; MF with posterior IT in notch of PT, no cleft under AT, MST indistinct to absent. Genital operculum as in fig. 60, p. 56 Muma (1951) with short, broad arms, long medial margin with a dark chitinized area midway, very slightly undu- late, wings short, posterior margin slightly curved (Fig. 55). No papillae on metatarsus of palpus; two tiny hairlike ctenidia were present on one specimen, the rest had none. BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 293 Female allotype: total length 19.0, chelicera length 4.6, chelicera width 1.8, propeltidium length 2.5, propeltidium width 3.6, palpus 13.0, first leg length 11.5, fourth leg length 21.5. Ratios: A/CP 6.60, GOL/GOW 0.75. Fe- male measurements (3): Total length 18.5- 21.0, chelicera length 4. 6-6. 7, chelicera width 2. 0-2.4, propeltidium length 2. 1-3.0, propel- tidium width 3. 2-4. 2, palpus length 11.5- 14.0, first leg length 10.5-13.0, fourth leg length 20.5-22.0. Ratios: A/CP 5.05-6.70, CL/CW 2.30-2.79, PL/PW 0.58-0.71, GOL/ GOW 0.72-0.82. Remarks. — Measurements were made from specimens collected at various times in Colorado National Monument and adjacent ar- eas. Four males but no females were found by Brookhart in pitfall traps during the summer of 1998. The range of collected specimens in- dicates an area within the central regions of the Colorado Plateau. Specimens examined. — UNITED STATES: Colorado: Mesa County, Colorado National Monument (39°03'N, 108°4rW), 15 July 1962, C. J. McCoy (d, CU); 21 June 1963, B. Vogel & C.J. McCoy (d, CU); 29 June 1973, C.J. McCoy (9, CU); 3 July 1973, C.J. McCoy (9, CU); Grand Junction (39°03'N, 108°33'W), 26 May-26 August 1998, Jack & Irene Brookhart (4 d, DMNS), Utah: Grand County, 32 Km west of Glade Park, coordinates unknown, 18 June 1951, no collection data (d, CU); Emery County, Castle Dale (39°12'N, llUOl'W), 26 June 1951, D.E. Beck (9, BYU); 7 July 1975, D. M. Allred (d, BYU); Sanpete County, Manti (39°16'N, 111°38'W), 21 June 1979, Ryan Olson (9, USU); Sevier County, Rich- field, (38°46'N, 112°05'W), 9 July 1963, G. F. Knowlton (9, BYU). Eremobates clarus Muma 1989 Figs. 33, 36, 45, 54 Eremobates clarus Muma 1989:10. Type . — Male holotype at 2194 meters in pitfall trap, Saratoga Stratton Experimental Watershed, Carbon County, Wyoming, US (4r27'N, 106°48'W), 17-21 July 1973, col- lected by John Schmid, AMNH. One male paratype from same trap (AMNH). Diagnosis. — Distinguished from closely re- lated E. scaber by less crimped, more smooth- ly curved EF in ectal view, and the cleft under the AT of the ME Female operculum has broader anterior arms and less undulation of the medial margin of the genital operculum. It is distinguished from E. ascopulatus by its pale coloration, slightly different female gen- ital operculum, and shape of ctenidia. Description. — Males.' Muma adequately described the male in 1989. Overall coloration very pale yellow, propeltidium dusky violet brown similar to E. scaber but with a larger median pale ovoid region, abdomen dusky, palpus and all legs pale yellow. Some speci- mens are duskier on propeltidium and ap- pendages. Thick, male FF gently curved to only slightly upturned in ectal view, no teeth, MF with posterior IT separate from PT, AT with anterior cleft, MST absent, FT graded I, III, II, IV, Fondal notch equal to or slightly wider (Fig, 33). Palpal papillae 33 — 80 +, two tiny, peg-like ctenidia (Fig. 36). Male holotype: Total length 19.0, chelicera length 4.7, chelicera width 2.3, propeltidium length 2.3, propeltidium width 3.4, palpus length 16.0, first leg length 12.0, fourth leg length 20.0. Ratios: A/CP 6.86, PL/PW 0.76, CL/CW 2.04, FL/FW 0.72. Male measure- ments(5): Length 18.0-22.0, chelicera length 4. 6-6. 6, chelicera width 2. 2-3. 2, propeltidium length 2. 2-2. 6, propeltidium width 3, 4-3. 6, palpus length 13.5-17.0, first leg length 11.0- 12.5, fourth leg length 17.0-21.0. Ratios: A/ CP 5.32-6.86, CL/CW 1.86-2.19, PL/PW 0.63-0.72, FL/FW 0.80-1.20, FW/FFW 2.'80- 3.75, CW/FFW 4.40-6.40. Eemales: Coloration as in males, chelicera typical of group. Posterior IT of MF separate. Cleft under AT, MST tiny. Genital operculum with short, broad arms, medial margin undu- late forming two small lobes, arms long, gent- ly curved, posterior margin rounded (Fig. 54). Female measurements(6).' Length 18.0- 24.0, chelicera length 4. 8-6.4, chelicera width 2.20-2.65, propeltidium length 1.6-2. 6, pro- peltidium width 3. 0-3. 6, palpus length 13.0- 15.5, first leg length 9.5-11.2, fourth leg length 16.0-21.0. Ratios: A/CP 4.92-6.97 CL/CW 2.04-2.50, PL/PW 0.50-0.72, GOL/ GOW 0.46-0.56. Remarks. — Eremobates clarus was collect- ed by Brookhart during the summer of 1998 from pitfall traps placed in both Pinon pine- Juniper and greasewood {Sarcobartus sp.) hab- itats in northwest Colorado near Dinosaur Na- tional Monument, It appears to occupy the area 294 THE JOURNAL OF ARACHNOLOGY Figures 27-35. — Fetal view male right chelicerae (arrow shows cleft under anterior tooth). 27. E. zinni. 28. E. similis. 29. E. mormonus. 30. E. actenidia. 31. E. scaber (arrow showing “crimp” in male fixed finger). 32. E. ctenidiellus. 33. E. clams. 34. E. hodai. 35. E. ascopulatus. Scale lines = 1 mm. BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 295 encompassed by the Laramie Plateau. The fe- male is described for the first time. Specimens examined. — UNITED STATES: Colorado: Moffat County, Castle Park, Dino- saur National Monument (40°28'N, 108°53'W), 18-30 June 1948, Hugo Rodeck (4 c?, CU); 4 July 1949, Hugo Rodeck (d, $, CU); 3.2 Km W of Craig (40°30'N, 107°32'W), 26 May-16 June 1971, Jack & Irene Brookhart (2 d’s, $, CU); 35.4 Km W of Craig, 18 June 1993, no collector ($, CU); 19.3 Km W of Craig, 1 May-26 July 1998, Jack & Irene Brookhart (3 (3, DMNS). Wyoming: Carbon County, Sara- toga Stratton Experimental Station (41°27'N, 106°48'W), Jon Schmid, (d, DMNS). Eremobates hodai Muma 1989 Figs. 17-19, 34, 40, 46 Eremobates hodai Muma 1989:13, fig. 14. Type . — Eremobates hodai male holotype labeled College of Idaho with no other infor- mation. Deposited in FSCA. Diagnosis. — Related to E. scaber and E. clarus from which it differs in coloration, shape of fondal notch, and female genital opercula. Although the holotype lacks palpal papillae and ctenidia other male specimens have papillae and ctenidia. Description. — Males.* Pale yellow colora- tion, palpus, legs pale, anterior region of pro- peltidium pale violet, eye tubercle dark. Che- licera pale with no stripes, typical dentition, FF crimped in ectal view, MF with posterior IT separate from PT, no cleft under AT (Fig. 34). Two short, pointed ctenidia (Fig. 40), 75- 106 palpal papillae (Fig. 19). Male holotype: Total length 21.0, chelicera length 6.5, chelicera width 3.3, propeltidium length 3.5, propeltidium width 4.0, palpus length 20.0, first leg length 17.5, fourth leg length 25.0. Ratios: A/CP 6.35, CL/CW 1.70, PL/PW 0.88, CL/CW 2.20, FL/FW 0.88, FW/ FFW 1.14, CW/FFW 4.00. Male measure- ments (3): Total length 17.0-23.0, chelicera length 6.25-6.46, chelicera width 2.17-3.00, propeltidium length 2.71-3.42, propeltidium width 3.54-4.67, palpus length 15.0-19.0, first leg length 11.0-15.5, fourth leg length 19.0-22.0. Ratios: A/CP 5.07-5.77, CL/CW 2.15-2.98, PL/PW 0.73-0.76, CL/CW 2.15- 2.98, FL/FW 0.95-1.00, FW/FFW 1.33-1.42, CW/FFW 4.73-5.17. Females: Coloration as in males. Chelicera typical, posterior IT separate from PT, no cleft under AT (Fig. 17). Genital opercula similar to E. scaber with long, broad arms, medial margin slightly curved inward, wings offset, long, posterior margin short ending in a point (Fig. 18). Female measurements (3): Total length 18.0-22.0, chelicera length 4.71-7.29, chelicera width 2.58-3.33, propeltidium length 2.83-3.33, propeltidium width 4.17- 5.00, palpus length 13.0-15.0, first leg length 11.0-13.0, fourth leg length 18.0-21.0. Ra- tios: A/CP 4.61-5.83, CL/CW 1.66-2.00, PL/ PW 0.66-0.70, GOL/GOW 0.82-0.86. Remarks, — Eremobates hodai was found in the Snake River Plain region of the Colum- bia River Plateau (USEPA 1986). Allred & Muma (1971) identified E. septentrionis, now E. ascopulatus and E. ctenidiellus from this region but were in error. Although this species may later be synonymized with E. clarus or E. scaber, its small A/CP and its pale lemon coloration leads us to separate it at this time. Specimens examined. — UNITED STATES: Idaho: Ada County, Eagle (43°4UN, 116°21'W), August 1987, W.H. & Mary Clark (2 9, ID); Butte County, 30 Km E of Arco, 6 June 1987, W.H. & Mary Clark {S , ID); 41.8 Km SE of Howe, W.H. & Mary Clark ( 9 , ID); 35.4 Km SE of Howe, 2-16 July 1987, P. Blom, W.H. Clark (d, ID). Oregon: Baker County, Sumpter (label says Idaho) (45°05'N, 1 13°44'W), 26 July 1995, W.H. & Mary Clark (d, 9, ID). Eremobates ascopulatus Muma 1951 Figs. 35, 43, 49, 53 Eremobates ascopulatus Muma 1951:60, fig. 19. Eremobates scaber (Kraepelin) sensu Muma 1951: 52-55, figs. 44-53 (not E. scaber Kraepelin). Eremobates septentrionis Muma 1970:12-14; Muma & Allred 1971:164; Brookhart 1972:33. NEW SYNONYMY. Eremobates flavus Muma 1989:11-12, figs. 7-9. NEW SYNONYMY Eremobates consors Muma 1989:11, figs. 5-6. NEW SYNONYMY Type. — Male holotype of E. septentrionis from Richfield, Utah, Sevier County, 20 June 1930, W.J. Gertsch (AMNH). Eremobates flavus: Male holotype collected in Reno, Washoe County, Nevada by W.E Hendrick, 7 July 1972; female allotype col- lected in Jungo, Humboldt County, Nevada by L.L. Stitt (holotype and allotype in FSCA). 296 THE JOURNAL OF ARACHNOLOGY 42 43 Figures 36-44. — Ventral view of male ctenidia. 36. E. clams. 37. E. mormonus. 38. E. ctenidiellus. 39. E. zinni. 40. E. hodai. 41. similes. 42. E. scaber. 43. E. ascopulatus. 44. Female E. similis. Scale lines = 1 mm. BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 297 Figures 45-51. — Dorsal view male propeltidia. 45. E. clarus. 46. E. hodai. 47. E. scaber. 48. E. cten- idiellus. 49. E. ascopulatus. 50. E. zinni. 51. E. mormonus. Scale lines = 1 mm. Eremobates consors: Female holotype from Minden, Douglas County, Nevada coU lected by D.A. Ball (Code #83H30-2 FSCA). Diagnosis. — Eremobates ascopulatus ap- pears to be related to E. clarus. It is distin- guished by the shape of male chelicera, col- oration of palpus and shape of female genital opercula. Description. — Male; Overall coloration dusky, straw yellow, chelicera with two dusky purple patches dorsally and one laterally, pro- peltidium tinged dusky purple anteriorly and laterally creating a broad, dusky yellow, ovoid 298 THE JOURNAL OF ARACHNOLOGY area (Fig. 49), abdomen grey to dark grey, palpal tarsus and metatarsus darker than the other appendages. Occasionally specimens have legs that are dusky at the tibia-femur joint. Some northern California specimens are darker, particularly the propeltidium and pal- pus. Male FF only slightly upturned in ectal view, posterior IT separate from PT, small, tri- angulate AT without a cleft, fondal notch LAV highly variable, MST medium (Fig. 35), 60- 80 + palpal papillae on most specimens, two short peg-like ctenidia (Fig. 43). Male holotype: Total length 21.0, chelicera length 5.76, chelicera width 2.73, propelti- dium length 3.1, propeltidium width 3.8, pal- pus length 18.0, first leg length 15.0, fourth leg length 23.5. Ratios: A/CP 6.60, CL/CW 2.10, PL/PW 0.81, FL/FW 1.20, FW/FFW 1.38, CW/FFW 6.20. Male measurements (S): Total length 19.5-23.0, chelicera length 5.25- 6.13, chelicera width 2.45-2.98, propeltidium length 2.80-2.98, propeltidium width 3.85- 4.20, palpus length 16.0-19.5, hrst leg length 14.5-16.5, fourth leg length 20.0-23.5. Ra- tios: A/CP 6.27-6.62, CL/CW 1.82-2.19, PL/ PW 0.68-0.73, FL/FW 0.74-1.22, FW/FFW 1.25-1.54, CW/FFW 4.86-7.08. Females: Coloration as in males. Chelicera typical of scaber group. MF with posterior IT in the notch of PT, no cleft anterior to AT. Genital opercula similar to E. ctenidiellus and E. geniculatus with long broad arms, slightly undulate medial margin, short wings, curved posterior margin (Fig. 53). Female measure- ments (4): Total length 18.5-27.0, chelicera length 5. 0-7. 6, chelicera width 2.4-3. 2, pro- peltidium length 2.4-3. 2, propeltidium width 3. 8-5.4, palpus 15.0-19.0, first leg length 10.0-14.5, fourth leg length 20.0-25.0. Ra- tios: A/CP 4.96-5.90, CL/CW 2.08-2.73, PL/ PW 0.55-0.74, GOL/GOW 0.42-0.80. Remarks. — Muma (1951) misidentified E. septentrionis as E. scaber but after examining the holotype of the latter in Europe erected E. septentrionis using his 1951 description of E. scaber (1970). He deviated from the original description in describing the palpal color as dusky purple. We here synonymize E. septen- trionis with E. ascopulatus based on exami- nation of both types which differ in only the absence of palpal papillae in E. ascopulatus. In this study we have found that in large sam- ples of papillate species there is an occasional specimen without papillae. Our ongoing re- search seems to indicate that papillae may arise sequentially after the penultimate molt. Eremobates ascopulatus is known from only two male specimens both of which were found in the same geographic area as E. septentrion- is. Because the description of E. septentrionis was based on specimens from many disparate parts of the western United States we have redescribed E. ascopulatus based on the ho- lotype and reinforced by sample specimens. Muma (1989) remarked as to the similarity between E. septentrionis and E. flavus and suggested that they may be the same species. We have found this to be the case. Muma’s (1989) holotype of E. flavus from Reno, Ne- vada had longer, thinner ctenidia but other specimens from this area and nearby north- eastern California had ctenidia and female genital opercula as in E. septentrionis. Some of the northern California specimens were darker in overall color suggesting perhaps a sibling species but more specimens are needed for examination. Eremobates consors is known from only one female and erected based on the shape of the female genital oper- culum. Examination of the holotype indicates that the operculum was desiccated and is in reality E. ascopulatus. Eremobates ascopulatus has a range that encompasses the Bonneville Basin and Lo- hantan Basin (see Trimble 1989) of northern Utah, Nevada, and northeastern California. It was also found at the Nevada Test Site (Muma 1963) where it is sympatric with E. zinni. Hall (1946) remarked as to the “intimate ecologi- cal connections” between the Lahontan basin and the Mojave Desert region of southern Ne- vada. These areas have the typical Great Basin desert shrub habitat. It appears to be related to E. clarus and E. ctenidiellus. Specimens examined. — UNITED STATES: California: Placer County, Lake Tahoe (39°10'N, 120°08'W), no date, Hubbard & Swartz, (d, FSCA); 11 July 1952, WJ. Gertsch ($, AMNH); Plumas County, Lake Almanor (40°13'N, 121°10'W), 7 July 1952, W. J. Gertsch (d, AMNH); 6.4 Km W of Quincy (39°56'N, 120°56'W), 21 June 1949, J. W MacSwain (3 6 AMNH); Shasta County, Cas- tella (41°08'N, 122°19'W), 26 July 1935, W J. Gertsch ($ , AMNH). Nevada: Elko County, 14.5 Km S of Contact, M.H. Muma (d, FSCA); Pershing County, Lovelock (40°10'N, BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 299 Figures 52-55. — Ventral view female genital opercula. 52. E. scaber. 53. E. ascopulatus. 54. E. clarus. 55. E. ctenidiellus. Scale line = 1 mm. 118°28'W), 22 June 1972, A.G. Rose ($, FSCA); Washoe County, Reno (39°3UN, 119°48'W), 15 August 1989, R C. Martenelli (2 cJ, $, FSCA). Utah: Box Elder County, Lucin (4r20'N, 113°54'W), 19 June 1952, D.E, Beck (2 d, BYU); Cache County, Logan (4r44'N, lir50'W), 4 July 1949, Steve Dewey (d, USU); 10 July 1995, D. Rasmus- sen (d, USU); Davis County, Hill Air Force Base (The county designation is probably in- correct as Hill Air Force Base Recreation Area is located in Weber County, not Davis County. See coordinates below under Weber County.), 17 July 1991, Mike Peterson (d, BYU); Salt Lake County, Sandy (40°35'N, lir53'W), 24 June 1985, Teresa Tipton (d, USU); Utah County, Alpine (40°27'N, lir46'W), 24 June 1997, A.L. Huillet (d, BYU); Environs, no coordinates, 14 June 1972, Troy Cooper ($, USU); Orem (40°17'N, 11L41'W), 20 June 1988, R. Wil- liams ($, BYU); 7 July 1993, Lisa Trotter (d, BYU); Provo (40°14'N, lir39'W), 1 Sep- tember 1993, D.O. White ($, BYU); Santa- quin (39°58'N, 111°47'W), 18 July 1987, J. Jarvis ($, USU); Spanish Fork (40°06'N, 1 1 r39'W), 8 July 1979, D. C. Holt ( $ , USU); Tooele County (says Uintah County on label), Vernon (40°05'N, 112°25'W), 7 August 1964, K. Bendixs ($, USU); Weber County, Hill Air Force Base (41°13'N, IIUSO'W), 27 July 1995, Larry Sanders (d, BYU). 300 THE JOURNAL OF ARACHNOLOGY Eremobates icenoglei new species Figs. 4-9 Type. — Male holotype collected by W. Icenogle in wet pit-fall trap, 29 August 1996, Winchester (33°42'N, 117°05'W), Riverside County, California; female allotype collected by W. Icenogle 22 Aug 1966 in wet pit-fall trap. Riverside County, California, 5 male par- atypes, 5 female paratypes collected by W. Icenogle, Winchester, California, Riverside County in wet pit-fall traps 29 August 1967- 23 August 1996. All types deposited in DMNS. Etymology. — Named for the collector Wendell Icenogle of Winchester, California. Diagnosis. — Eremobates icenoglei new species appears most closely related to E. zin- ni but is separated from it and others of the scaber group by absence of AT on male MF, a fond that is noticeably longer than wide, and a thickened FF. The female genital operculum is distinctive (Fig. 9). Description. — Males.- Coloration overall dark to dusky yellow, abdominal tergites dusky, appendages dusky yellow with palpal metatarsus and the tibia-femora joint area dusky violet-brown, propeltidium dusky pur- ple on anterior edge and top lateral one third (Fig 4). Fixed finger of chelicera with little or no crimping, fondal notch longer than wide; width of FF 80% the width of FN; MF with large primary tooth, and a ridge that is slightly elevated anteriorly instead of intermediate and anterior teeth, MST intermediate in size (Fig. 5-6). Four stiletto shaped ctenidia on first post-spiracular sternite extending approxi- mately half the length of the sternite (Fig. 7); no palpal papillae. Male holotype: Total length 19.0, chelicera length 4.8, chelicera width 2.2, propeltidium length 2.6, propeltidium width 3.5, palpus length 16.0, first leg length 16.0, fourth leg length 23.0. Ratios: A/CP 6.82, CL/CW 2.20, FL/FW 1.40, WFF/FW 1.20, CW/WFF 4.23. Male paratypes (5): Total length 18.0-24.0, chelicera length 5.50-6.92, chelicera width 4. 4-6.4, propeltidium length 2.32-3.20, pro- peltidium width 3. 2-4.0, palpus length 14.0- 17.0, first leg length 10.0-14.0, fourth leg length 17.0-24.0. Ratios: A/CP 5.52-6.80, CL/CW 1.73-2.20, FL/FW 1.17-1.44, FW/ FFW 2.29-3.75, CW/WFF 4.23-5.75. Eemale: Coloration the same as male. Che- licera typical of species; MF with posterior IT separate from PT, MST indistinct to absent (Fig. 8), no papillae on metatarsus of palpus; 2-3 tiny hairlike ctenidia were present on two specimens, the rest had none. Genital oper- culum with longer, broad arms, a long, slightly recurved medial surface ending in a point, wing short to absent, and posterior edge trun- cated (Fig. 9). Female (allotype): Total length 19.0, chelicera length 6.2, chelicera width 2.8, propeltidium length 2.0, propeltidium width 3.6, palpus length 12.0, first leg length 9.0, fourth leg length 17.5. Ratios: A/CP 4.70, CL/ CW 2.20, PL/PW 0.55, GOL/GOW 0.82. Fe- male paratypes (5): Total length 17.0-26.0, chelicera length 4. 8-7. 6, chelicera width 1.8- 3.4, propeltidium length 2, 0-3.0, propeltidium width 3. 5-5.0, palpus length 12.0-16.0, first leg length 9.0-13.0, fourth leg length 16.0- 22.0. Ratios: A/CP 4.70-5.66. CL/CW 2.20- 2.67, PL/PW 0.48-0.63, GOL/GOW 0.74- 0.82. Remarks. — Eremobates icenoglei appears to be restricted to the Coastal Chaparral hab- itat. It was found by Wendell Icenogle in buildings and in pitfall traps. Specimens examined. — UNITED STATES: California: Riverside County, Winchester (33°42'N, 1I7°05'W), 29 August 1967 (d, 3 9), 16 September 1967 (2 9); 24 September 1968 (9), 11 August 1973 (3 d), 3 September 1973 (3 9), 4 September 1973 (9), 8 August 1981 (4 d), 17 August 1987 (3 d), 16 August 1988 (2 d), 1 August 1996 (2 d), 23 August 1996 (d). All collected by Wendell Icenogle. Types and paratypes in DMNH. Eremobates mormonus (Roewer 1934) Figs. 29, 37, 51, 57 Eremoperna mormona Roewer 1934:561, figs. 323e, 324f. Eremoperna mormonus (Roewer); Muma 1962:1; Muma 1970:12, fig. 9; Brookhart 1972:32. Eremobates geniculatus (Simon) sensu Muma 1951:55-57 (misidentification). Eremobates mimbreniis Muma 1989:12-13, figs. 10-13. NEW SYNONYMY. Type . — Female holotype of Eremoperna mormona from Utah, SMF/RII/3446. No spe- cific locality or collector indicated. Male ho- lotype of E. mimbrenus from Signal Peak (32°55'N, 108°10'W), Grant County, New Mexico, 17 June 1976 and female allotype BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 301 60 Figures 56-60. — Ventral view female genital opercula. 56. E. similis. 57. E. mormonus. 58. E. zinni. 59. E. actenidia. 60. E. legalis. Scale lines = 1 mm. from same locality collected on 1 July 1976. Both collected by Martin Muma (FSCA), Diagnosis. — This species appears closely related to E. similis Muma but can be distin- guished from it by its darker color, shape of fondal notch, presence of papillae, and thick- ness and length of ctenidia. Description. — Males: Coloration dusky straw yellow over all, propeltidium dusky pur- ple anteriorly and on the lateral one third (Fig. 51), tergites with broad violet brown stripe, abdomen grey, palpal tarsus and metatarsus dusky amber, legs dusky amber on femur-tibia joint. FF regularly curved; MF with large PT and AT, no cleft under AT, posterior IT in the notch of PT, MST tiny to absent. Fondal notch wider than long (Fig. 29); ctendia 4 flat, sword like, extending slightly more than half the length of succeeding sternite (Fig. 37); 80+ papillae on palpal scopula. Apical plumose 302 THE JOURNAL OF ARACHNOLOGY bristle covers most of mesal groove. Male measurements (5): Total length 15.0.-23.0, palpus length 10.0-22.0, first leg length 9.0- 14.5, fourth leg length 14.5-23.0, chelicera length 3. 5-5. 4, chelicera width 1.6-2. 6, pro= peltidium length 1.6-2. 8, propeltidium width 2.00- 3.85. Ratios: A/CP 6.08-7.59, CL/CW 1.94-2.20, PL/PW 0.70-0.80, FL/FW 0.50- 0.67, FW/FFW 1.25-1.88, CW/FFW 4.60- 6.20. Females: Coloration same as males. Che- licera typical of species; MF with posterior IT in the notch of PT, no cleft under AT. MST indistinct to absent. Genital operculum similar to E. zinni and E. socal, new species as in fig. 68, pg. 59, Muma 1951 with short arms, me- dial surface gently curved ending in a point as in E. zinni and E. similis, short curved wing (Fig. 57). No papillae on metatarsus of palpus; seven tiny hairlike ctenidia were present on one specimen. Eemale allotype: Total length 19.0, chelic- era length 5.1, chelicera width 2.2, propelti- dium length 2.3, propeltidium width 3.9, pal- pus length 12.5, hrst leg length 11.0, fourth leg length 18.5. Ratios: A/CP 5.75, CL/CW 2.31, PL/PW 5.90, GOL/GOW 0.54. Eemale measurements (3): Total length 18.5-21.0, chelicera length 4.40-5.02, chelicera width 2.0- 2. 1, propeltidium length 2. 1-3.0, propel- tidium width 3.4-3. 6, palpus length 11.5- 12.0, first leg length 10.5-12.0, fourth leg length 18.0-22.0. Ratios: A/CP 5.34-6.16, CL/CW 2.20-2.40, PL/PW 0.58-0.94, GOL/ GOW 0.54-0.63. Remarks. — This species was erroneously described as E. geniculatus by Muma 1951, but after examining the holotype in Paris he identihed it as Roewer’s species from Utah. Muma (pers. comm.) stated that the “type” of E. mormonus in the collection of MNHN is not Koch’s type but may be a lectotype set up by Roewer and therefore invalid. No other lo- cality data were given but it must have been somewhere in southwest Utah based on the presence of other species in that area. We found E. mormonus in northern Arizona around the Grand Canyon, SW Colorado and western New Mexico on mesas with sage, pi- hon pine-juniper and alpine meadows. The Arizona specimens are darker in overall col- oration. We here synonymize E. mimbrenus with E. mormonus. Eremobates mimbrenus was described by Muma (1989), and the de- scription that Muma gave is correct except that in his drawing of the right male chelicera, the IT is shown on the PT yet on the type specimen itself, the IT is separate (Muma 1989, fig. 12). In the same vial is a male and female, collected by Muma that appear to be E. mormonus as well as two other vials from the area with two males and a female which also key out to E. mormonus. Muma’s (1989) description of the female allotype of E. mim- brenous was based on a female with three in- termediate teeth on the movable finger col- lected in a pitfall trap in SW New Mexico. A female in the same vial does not have this character and could be recognized as E. mor- monus based on genital opercula and chelicera structure. The male in the same vial is also recognized as E. mormonus. Until more spec- imens are collected from this area we have synonymized E. mormonus and E. membren- us. Specimens examined. — UNITED STATES: Arizona: Coconino County, North rim of Grand Canyon (36°12'N, 112°03'W), 13 July 1934, Rockefeller, (2 9, AMNH); 18 July 1934, Lutz (d, 9, AMNH); 13 June 1934, Wilton Ivie (d, AMNH); Flagstaff (35°1 UN, 1 1 1°39'W), 22 July 1949, Billy Hughes (9, AMNH); Kaibob Forest (35°50'N, 112°05'W), 14 June 1934, Ivie & Rasmussen (d, AMNH). Colorado: Montezuma County, Chimney Rock (37°04'N, 108°43'W), 31 July 1973, B. Vogel (2 d’s, CU); McFee Reservoir (misspelling of Mc- Phee Reservoir) (37°34'N, 108°34'W), 28 Au- gust 1997, B. Jacobi (2 d, 2 9, CSU); Miller Reservoir (37°37'N, 108°27'W), 28 August 1997, no collector (d, CSU); Mesa Verde Na- tional Monument (37°14'N, 108°28'W), 23 July 1941, no collector (2 d, 9, CU). New Mexico: Grant County, Signal Peak, Gila Na- tional Forest (32°55'N, 108°10'W), 6 July 1976, 1 July 1976), Martin Muma (2 d, 9, FSCA). Eremobates similis Muma 1951 Figs. 28, 41, 56 Eremobates similis Muma 1951:figs. 70—71; Muma 1962:4; Muma 1970:14; Muma 1989:9. Type. — Male holotype. Elk Ridge, Utah County, Utah (40°00'N, 11U40'W), 13 June 1936, Douglas Henriques, originally deposited at University of Utah but now at AMNH. Diagnosis. — Lighter coloration with a more violet tinge than the closely related E. mor- BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 303 monus with 4-6 short, thin, needle like cte- nidia extending less than half the length of succeeding sternite, no palpal papillae. Description. — Males: Appendages and propeltidium straw yellow with violet brown markings. Propeltidium lightly blotched violet brown on anterior and lateral margins, palpus and legs violet brown on femur, tibia, meta- tarsus, tarsus, abdominal tergites yellow grey. FF regularly curved in ectal view with no teeth; MF with posterior IT situated in the notch; AT triangulate with no cleft; MST tiny to absent. Fond significantly wider than long (Fig. 28); 4-6 thin, needle like ctenidia on first post spiracular sternite extending less than half the length of the sternite (Fig. 41). No papillae on palpal scopula. Both male and female descriptions are made from specimens collected in wet pit fall traps in Costillo Coun- ty, Colorado in the summer of 1997 by J.O. and I.P. Brookhart. They were the most com- mon solifugid found in the San Luis Valley of Colorado. They were referred to as E. mor- monus by Brookhart (1972). Male holotype: Total length 22.0, chelicera length 5.2, chelicera width 2.6, propeltidium length 2.5, propeltidium width 3.9, palpus length 17.0, first leg length 14.0, fourth leg length 22.0. Ratios: A/CP 6.88, CL/CW 2.00, PL/PW 0.64, FL/FW 0.50, FW/FFW 1.66. Male measurements (6) (from San Luis Valley, Colorado): Total length 15.5-17.0, chelicera length 4. 2-4. 8, chelicera width 1. 8-2.0, pro- peltidium length 2. 0-2. 2, propeltidium width 2. 8-3. 2, palpus 12,0-18.0, first leg length 11.0- 16.5, fourth leg length 17.0-20.0. Ra- tios: A/CP 6.52-1 m, CL/CW 2.22-2.44, PL/ PW 0.63-0.71, FL/FW 0.62-0.71, FW/FFW 3.50-4.33, CW/FFW 5.50—6.50. Females: Coloration as in males, chelicera typical of species; MF with posterior IT in the notch of PT; MST indistinct to absent. Genital operculum with short, thin arms, interior lat- eral surface curved ending in a point as in E. zinni and E. mormonus, long recurved wings, rounded posterior margin (Fig. 56). No papil- lae on metatarsus of palpus; 2-5 tiny hairlike ctenidia were present on most specimens (Fig. 44). Female measurements (6): Total length 16.5-22.0, chelicera length 4.20-5.52, chelic- era width L8-2.4, propeltidium length 2.0- 2.6, propeltidium width 3. 0-4.0, palpus length 10.0- 15.0, first leg length 8.0-11.5, fourth leg length 14.0-21.0. Ratios: A/CP 5.30-5.76, CL/CW 2.33-2.88, PL/PW 0.62-0.81,, GOL/ GOW 0.58-0.71. Remarks. — Eremobates similis was de- scribed from a specimen with a locality label from Salt Lake City, Utah. Although the type locality for E. similis is listed as Salt Lake City, Utah we have collected this species only in an area that can be roughly called the north- ern Rio Grande Valley at three sites, San Luis Valley, Colorado, Seviletta LTER, Socorro County, New Mexico, and Bandelier National Monument, Sandoval County, New Mexico. Examination of the type fits Muma’s descrip- tion but we were unable to determine if there was an error in labeling the type locality. None of the specimens from Utah that we used in the scaber study could be identified as E. similis. Females of this species have not been previously described. Each of the above populations occurred in high desert shrub habitat and varied somewhat in color. The male population of Bandelier National Monument had a statistically signif- icant variation in A/CP ratio indicating the possibility of sibling species. San Luis Valley (2303 m) specimens were collected in Rabbit Bush, snake weed, greasewood habitats. Ban- delier National Monument specimens from Pi- non- Juniper and Ponderosa Pine habitat and Sevilleta LTER from Pinon-Juniper habitat but not from grassland and creosote bush (Brookhart & Brantley 2000). Specimens examined. — UNITED STATES: Colorado: Costillo & Saguache Counties, San Luis Valley, 8 June-8 August 1997, Jack & Ir- ene Brookhart in wet pitfalls (22 (3, 19 9); New Mexico: Socorro County, Sevilleta National Wildlife Refuge (34°03'N, 106°23'W), 1989- 1994 in wet pitfalls (7 (3, 18 9); Sandoval County, Bandelier National Monument (35°47'N, 106°18'W), 1998-2000 in wet pit- fall traps (22 (3, 19 9). Deposited at DMNS and University of New Mexico. Eremobates actenidia Muma 1989 Figs. 30, 59 Eremobates actenidia Muma 1989: 9-10, figs. 1, 2. Type . — Male holotype from Gouldings Trading Post, Monument Valley, San Juan County, Utah, USA (37°06'N, llOHl'W), 2 June 1953, R.E. Ryckman, R.D. Lee, C.T Ames, C.C. Lindt, C.T Christianson. Depos- ited AMNH. 304 THE JOURNAL OF ARACHNOLOGY Diagnosis. — This specimen may be sepa- rated from all but E. ctenidiellus by its lack of ctenidia on the first post stigmatal segment. Eremobates ctenidiellus generally lacks cte- nidia but can easily be distinguished by its pale coloration as opposed to the dark color- ation of E. actenidia and the shape of the fon- dal notch, which is significantly longer than wide. It has a high A/CP ratio indicating lon- ger appendages and a statistically thinner male fixed finger in relation to the fondal notch. Description. — Males: Appendage and pro- peltidium coloration dusky yellow to brown- ish yellow, propeltidium tinged brownish vi- olet on anterior and lateral margins, palpus tinged brownish violet on tarsus and metatar- sus, legs dusky yellow. Cheliceral FF regular- ly curved, MF with small triangulate AT, no cleft, small IT with the posterior separate from the PT (Fig. 32), no ctenidia, 70 + palpal pa- pillae. Male holotype: Total length 18.0, che- licera length 4.0, chelicera width 2.3, propel- tidium length 2.3, propeltidium width 3.5, palpus length 17.0, first leg length 13.0, fourth leg length 20.0. Ratios: A/CP 7.94, CL/CW 1.74. PL/PW 0.66, FL/FW 1.02, FW/FFW 1.70, FW/CW 6.09. Male measurements (4): Total length 17.50-21.00, chelicera length 4.92-6.25, chelicera width 2.29-2.79, propel- tidium length 2.17-2.83, propeltidium width 3.75-4.58, palpus length 16.5-22.0, first leg length 12.0-13.0, fourth leg length 19.5-23.0. Ratios: A/CP 5.94-6.78, CL/CW 2-2.42, PL/ PW 0.58-0.68, FL/FW 0.79-1.06, FW/FFW 1.56-2.00, CW/FFW 6.09-8.13. Females: Coloration as in male except the legs are lightly tinged violet at the tibia femur joint. Chelicera typical, MF with large PT, 2 IT, large AT, posterior IT in notch of PT, no cleft under AT, tiny to absent MST Genital opercula with long, thin arms, a slightly curved interior margin ending in a lobe, wings offset, posterior margin truncate (Fig. 59). Female measurements (5): Total length 16.0-20.5, chelicera length 4. 8-5. 8, chelicera width L8-2.4, propeltidium length 2. 2-2. 8, propeltidium width 3. 2-4.4, palpus length 14.5-16.0, first leg length 9.5-1 1.0, fourth leg length 16.0-17.0. Ratios: A/CP 4.88-6.37, CL/CW 2.00-2.67, PL/PW 0.57-0.75, GOL/ GOW 0.67-0.80. Remarks. — Eremobates actenidia has only been found in the desert grass region of San Juan County, Utah. Brookhart collected from pitfall traps set in three different habitats along alb km stretch of Hwy 195 in San Juan County, Utah from 29 May 2000-28 August 2000 and again on 6 June 2001. Eremobates actenidia was collected from desert grasslands but not from desert shrub or Pinon- Juniper as- semblages in this transect. Eremobates mor- monus is found 161 km east in Montezuma County, Colorado and E. corpink, new spe- cies, is found 161 km west in the Coral Pink Sand Dunes of Kane County, Utah at approx- imately the same latitude. Specimens examined. — UNITED STATES: Utah: San Juan County, 6.4 Km N of Bluff (37°17'N, 109°33'W), 10 June-26 August 2000, Jack & Irene Brookhart in wet pitfall traps (3 6, 4 9, DMNS). Eremobates socal new species Figs. 10-16 Types. — Male holotype, female allotype; California, San Diego County, Mt. Palomar St. Park (33°20'N, 116°54'W), 13 July 1953, W.J. & J.W. Gertsch (AMNH). Etymology. — A noun in apposition refer- ring to the type locality, Southern California, as used by Jim Rome, radio and TV sports talk show host. Diagnosis. — Eremobates socal can be sep- arated from E. zinni and E. mormonus by size and shape of ctenidia, color variation and fe- male genital operculum. Description. — Males: Appendage and pro- peltidium background coloration dusky yel- low, propeltidium violet brown on the anterior and lateral fringes, a broadly ovoid yellow center (Fig. 10), abdomen dusky grey, apical half of palpal metatarsus dusky violet, legs I & II dusky yellow, femur of legs III & IV dusky violet. Male cheliceral FF regularly curved, MF with large PT, small triangulate AT with a cleft, small IT, posterior IT separate from PT (Figs. 11-12), 47-60 palpal papillae, a few on tarsus (Fig. 13), 4 short needle like ctenidia extending less than half the succeed- ing segment (Fig. 14). Male holotype: Total length 19.0, chelicera length 5.63, chelicera width 2.58, propelti- dium length 2.50, propeltidium width 4.36, palpus length 18.0, first leg length 13.0, fourth leg length 21.5. Ratios: A/CP 6.46, CL/CW 2.18, PL/PW 0.57, FL/FW LOO, FW/FFW 1.45, CW/FFW 5.64. Male paratypes (4): To- tal length 19.0-20.0, chelicera length 5.00- BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 305 5o63, chelicera width 2.42—2.58, propeltidium length 2.50-2.92, propeltidium width 4.17- 4.38, palpus length 16.0-18.0, first leg length 12.0-14.0, fourth leg length 20.0-21.5. Ra- tios: A /CP 6.06-6.46, CL/CW 2-2.18, PL/ PW 0.62-0.7. FL/FW 0.71-1.00, FW/FFW 1.17-1.70, CW/FFW 5.00-6.00. Females: Coloration as in the male. Chelic- erae typical with posterior IT of MF in the notch, no cleft under AT (Fig. 15). No palpal papillae, no ctenidia. Genital operculum with long, broad arms, long curved medial edge ending in a point, short, offset wings, curved posterior margin (Fig. 16). Female allotype: Total length 19.0, chelic- era length 5.29, chelicera width 2.08, propel- tidium length 2.29, propeltidium width 3.83, palpus length 12.0, first leg 12.0, fourth leg length 19.0. Ratios: A/CP 5.27, PL/PW 0.60, CL/CW 2.54, GOL/GOW 0.72. Female par- atypes (2): Total length 20.0-20.5, chelicera length 6.25-6.50, chelicera width 2.5, propel- tidium length 2.5 — 2.6, propeltidium width 4.50-4.58, palpus length 14.5-15.0, first leg length 12.0, fourth leg length 18.0-19.0. Ra- tios: A/CP 5.97-6.00, CL/CW 2.5, PL/PW 0.57, GOL/GOW 0.5-0.6. Remarks. — Eremobates socal is similar to E. zinni and E. mormonus but can be separat- ed by the shape and size of the ctenidia, col- oration, and shape of female genital opercula. It was found in the coastal shrub area on Mt. Palomar in San Diego County, California. Specimens examined. — UNITED STATES: California: San Bernardino County, Big Bear Lake (34°14'N, 116°54'W), July 1950, no collector (6, AMNH); Granite Cove, 8 Km N on Kelbaker Road (34°46'N, 115°39'W), E. Faster (2 S , California State Riverside); Josh- ua Tree National Monument (35°07'N, 116°02'W), 22 August 1994, no collector (9, DMNH); San Diego County, Mount Palomar State Park (now Palomar Mountain State Park) (33°20'N, 116°54'W), 13 July 1953, W.J. & J.W Gertsch (3 6,9, AMNH). Eremobates zinni Muma 1951 Figs. 27, 39, 50, 58 Muma 1951:58, figs. 65-68; Muma 1970:14; Muma 1987:1920. Type . — Male holotype, female allotype from Las Vegas, Clark County (36°10'N, 115°08'W), Nevada in AMNH. Collected by Donald J. Zinn, May through August 1944. Female paratype from Las Vegas, Clark Coun- ty (36°10'N, 115°08'W), Nevada, February through June 1945 (AMNH). Etymology. — Named for the collector Don- ald Zinn. Diagnosis. — Eremobates zinni is closely re- lated to E. socal new species. It can be sep- arated by the shape of the male chelicera, col- oration, and shape of female genital opercula. Description. — Males.- Appendages dusky yellow with dusky violet brown on tarsus and distal end of metatarsus of palp, propeltidium dusky yellow with light violet tinges on an- terior and anterio-lateral edges (Fig. 50). Male cheliceral FF regularly curved, MF with small triangulate AT with cleft, posterior IT separate from PT; fondal notch longer than wide (Fig. 27). Ctenidia 4 short, flat (Fig. 39), 40-80 pal- pal papillae. Male holotype: Total length 21.0, chelicera length 5.7, chelicera width 2.5, propeltidium length 3.0, propeltidium width 6.4, palpus length 19.0, first leg length 15.0, fourth leg length 23.0. Ratios: A/CP 6.55, CL/CW 2.28, PL/PW 0.47, FL/FW 1.17, FFW/FW 1.33, CW/FFW 0.80. Male measurements (1): Total length 14.0, propeltidium length 3.12, propel- tidium width 6.4, chelicera length 6.24, che- licera width 3.0, palpus length 12.0, first leg length 10.0, fourth leg length 16.0. Ratios: A/ CP 4.10, CL/CW 2.08, PL/PW 0.49, FL/FW 0.90, FFW/FW 1.38, CW/FFW 1.10. Females: Same coloration as males, MF with posterior IT in the notch of PT, AT with no cleft, genital opercula with long, thin arms, a curved medial edge ending in a point, short offset wings and a curved posterior surface (Fig. 58). Female allotype: Total length 23.0, propeltidium length 3.9, propeltidium width 7.0, chelicera length 4.8, chelicera width 2.5, palpus length 15.0, first leg length 13.0, fourth leg length 21.0. Ratios: A/CP 4.80, CL/CW 2.52, PL/PW 0.56, GOL/GOW 0.68. Remarks. — Eremobates zinni and E. as- copulatus are sympatric at the Nevada Test Site. Eremobates zinni appears to be related to both E. socal new species and E. corpink new species. Specimens examined. — UNITED STATES: Nevada: Clark County, Las Vegas (36°10'N, 115°08'W), May-August 1944, Donald J. Zinn (d, AMNH). Utah: Washington County, Saint George (37°06'N, 1 13°35'W), no date, no col- lector (6, BYU). 306 THE JOURNAL OF ARACHNOLOGY Eremobates corpink new species Figs. 20-26 Type. — Male holotype from Kane County, Utah, Coral Pink Sand Dunes (37°05'N, 112°40'W), 2 August 1988, R.W. Bauman (DMNH). Female allotype from Kane County, Utah, Coral Pink Sand Dunes, 1 1 June 2002, S.M. Clark (DMNS). One male paratype from Kane County, Utah, Escalante National Mon- ument (37°25'N, 1 1 1°33'W), 9 July 2000, DJ. Craven, E. Cygen and W.N. Wendel (BYU). One male from Kane County, Utah, Coral Pink Sand Dunes, 9 July 2002, S.H. Clark and J.S. Robertson (BYU). Diagnosis. — Separated from E. zinni and E. ascopulatus by size and shape of ctenidia, shape of female opercula and coloration. Etymology. — A noun in apposition refer- ring to the Coral Pink Sand Dunes, Kane County, Utah. Description. — Males.- Overall coloration pale straw yellow, palpal tarsus and distal one third of metatarsus blotched brownish violet, all other appendages pale straw yellow, pro- peltidium with lightly blotched brownish vi- olet except for pale median ovoid region (Fig. 20), abdomen yellow to grey. Cheliceral fixed finger thin, regularly curved, movable finger with large PT, smaller triangulate AT, cleft an- terior to AT, posterior IT separate from PT (Figs. 21-22). Two short, thin ctenidia (Fig. 24) , palpal papillae 40-82 (Fig. 23). Male ho- lotype: Total length 21.0, propeltidium length 2.5, propeltidium width 2.9, chelicera length 4.7, chelicera width 2.4, palpus length 16.0, first leg length 12.5, fourth leg length 22.0. Ratios: A/CP 6.92, PL/PW 0.90, CL/CW 1.96, FL/FW 0.56, FFW/FW 1.80, CW/FFW 12.00. Male paratypes (2): Total length 20.0- 20.5, propeltidium length 2.60-3.82, propel- tidium width 3.15-3.90, chelicera length 3.82-5.45, chelicera width 2. 0-3. 9, palpus length 16.0-17.5, first leg length 12.0-14.0, fourth leg length 22.0-22.5. Ratios: A/CP 6.22-6.94, PL/PW 0.80-0.98, CL/CW 1.32- 2.60, FFW/FW 1.19-1.45, FL/FW 0.56-0.94, CW/FFW 7.96-10.70. Female allotype: Overall coloration as in male, palpus and legs yellow, propeltidium blotched violet purple in anterior edge, abdo- men grey, chelicera typical, no cleft under AT of MF, posterior IT in notch of PT of MF (Fig. 25) , 2 faint violet stripes on posterior dorsal edge of chelicera, genital opercula with short, broad arms, slightly curved medial edge with two small lobes, wings short and curved, pos- terior edge curved (Fig. 26). Female allotype: Total length 19.0, propeltidium length 2.5, propeltidium width 2.8, chelicera length 6.0, chelicera width 2.8, palpus length 15.0, first leg length 12.0, fourth leg length 20.0. Ratios: A/CP 5.53, CL/CW 2.15, PL/PW 0.90, GOL/ GOW 0.73. Remarks. — Eremobates corpink appears related to E. zinni. The restricted habitat in and around the Coral Pink Sand Dunes has produced other endemic species (Knisley & Hill 2001). Specimens examined. — UNITED STATES: Utah: Kane County, Escalante National Monu- ment (37°25'N, 11U33'W), 9 July 2000, D.J. Craven, E. Cygen, & W.N. Wendel (d para- type, BYU); Coral Pink Sand Dunes (37°02'N, 1 12°42'W), 9 July 2002, S.H. Clark & J.S. Robertson (d paratype, BYU). Eremobates legal is Harvey 2002 Pig. 60 Datames geuiculatus (C.L. Koch) Simon 1879:138. (Not E. geuiculatus Simon, sensii Muma 1951). Eremocosta geuiculata (Simon) Roewer 1934:570. Eremobates legalis Harvey 2002:451. Type. — Female holotype from Mexico, No. 2129 (Roewer No. 9135) in MNHN Etymology. — From the Latin “legalis” which means according to the law (Harvey 2002). The original specific epithet of “geni- culatLis” is from the Latin meaning bent. This origin is interesting since it probably refers to the male fixed finger and E. geuiculatus is known only from one female. Diagnosis. — Muma (1970) distinguishes this species by the presence of two small cte- nidia but this is invalid since females of sev- eral species possess ctenidia. Female genital operculum is similar to E. ascopulatus. Description. — Female: Overall coloration pale yellow. Palpus, legs yellow, propeltidium with faint violet tinge anteriorly. Abdomen grey. Chelicera typical, posterior tooth of MF in the notch of PT, no cleft. No palpal papillae, two tiny hair like ctenidia. Operculum with long, thin arms, smooth medial margin, short gently curving wings, posterior margin curved (Fig. 60). Female holotype: Total length 23.0, chelicera length 5.4, chelicera width 2.5, pro- peltidium length 2.3, propeltidium width 4.3, BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 307 palpus length 13.0, first leg length 10.5, fourth leg length 14.5. Ratios: A/CP 4.93, CL/CW 2.16, PL/PW 0.53, GOL/GOW 0.69. Remarks. — Simon's (1879) description of Datames geniculatus was of a specimen dif- fering from Roewer’s 1934 description of Er- emocosta geniculata. Harvey clarified this in 2002 by designating the type (Roewer No. 9135) Eremobates legalis. Eremobates legal- is/E. geniculatus is known from the type only. Muma (pers. comm.) indicates that the other specimen in the vial is an immature. It is the only member of the scaber group presently known from Mexico (Vasquez 1981; Muma 1987). We have examined other specimens from the southwestern United States labeled E. geniculatus by Muma but now consider them misidentifications. KEY TO THE MALES OF EREMOBATES SCABER GROUP (Females can best be distinguished by the shape of genital opercula) 1. Ctenidia none or hair-like 2 Ctenidia present 3 2. Ctenidia absent, palpal tarsus, metatarsus brownish violet .......... Eremobates actenidia Ctenida absent or 2 hair-like (Fig. 38), palpal tarsus and metatarsus pale .......... . Eremobates ctenidiellus 3. Two ctenidia present 4 More than two ctenidia present .............................................. 8 4. Fixed finger severely crimped in ectal view (Fig. 1); ctenidia short, thin, pointed (Fig. 42); palpus dusky straw yellow Eremobates scaber Fixed finger with no or slight crimping in ectal view; ctenidia variable; palpus variable 5 5. Overall coloration lemon yellow; ctenidia flat, sword-shaped (Fig. 40) . . . Eremobates hodai Overall coloration dusky yellow, brown or brownish violet; ctenidia not sword-shaped . . 6 6. Palpal color dusky yellow; posterior intermediate tooth separate from principle tooth of movable finger; no cleft under anterior tooth of movable finger; fond length and width equal (Fig. 35); ctenidia flat, needle-like (Fig. 43) Eremobates ascopulatus Palpal color dusky yellow to brownish violet; posterior intermediate tooth in the notch of primary tooth of movable finger; cleft under anterior tooth of movable finger; fond length to width ratio equal or wider; ctenidia variable 7 7. Palpal color dusky yellow; cleft under anterior tooth of movable finger; fond length to width ratio equal (Fig. 33); ctenidia broad, flat (Fig. 36) .............. Eremobates clarus Palpal coloration brown violet on tarsus and metatarsus; cleft under anterior tooth of mov- able finger; fondal notch wider than long; ctenidia short, thin Eremobates corpink 8. Anterior tooth of movable finger absent or an undifferentiated ridge; fondal notch longer than wide (Fig 5); four flat ctenidia (Fig. 7) Eremobates icenoglei Anterior tooth of movable finger present; fondal notch ratio equal or wider; three or four ctenidia, thin or flat ........................................................ 9 9. Ctenidia flat 10 Ctenidia thick, needle-like . 11 10. Three to four ctenidia present, stiletto-like, extending half the length of succeeding sternite (Fig. 39); palpal tarsus and distal end of metatarsus dusky; posterior intermediate tooth of movable finger separate from principle tooth; cleft under anterior tooth of movable finger (Fig. 27) ..................................................... Eremobates zinni Four, flat ctenidia present (Fig. 37); palpal tarsus and metatarsus dusky amber; posterior intermediate tooth of movable finger in notch of primary tooth; no cleft under anterior tooth of movable finger (Fig. 29) ............................. Eremobates mormonus 11. Four to six thin, short, hair-like ctenidia present (Fig. 41); no palpal papillae; palpus brown- ish violet on tarsus, metatarsus, tibia, and tib-fib joint; posterior intermediate tooth of movable finger in the notch of principle tooth; no cleft under anterior tooth (Fig. 28) . . . ............................................................ Eremobates similis Four, flat ctenidia present (Fig. 14); papillae present (Fig. 13); palpus dusky brown on 308 THE JOURNAL OF ARACHNOLOGY tarsus and metatarsus; posterior tooth separate from principle tooth of movable hnger; cleft under anterior tooth of movable finger (Fig. 11) Eremobates socal CLADISTIC ANALYSIS To our knowledge, no previous cladistic analysis has been attempted for any genus or species group of Solifugae. Herein we present the first testable hypothesis of the relation- ships among species in the E. scaber species group (Fig. 61) with a member of the Ere- mobates pallipes species group, E. palUpes (Say 1823) set as the outgroup based on the simple structure of male chelicera and female genital opercula. Many of the ratios that are useful as diagnostic characters, are too vari- able to be useful as characters for cladistic analysis. We limited our cladistic analysis to characters that could be coded objectively. The following 12 characters were used to analyze the relationships among the species in the E, scaber species-group (see Table 1). 1 ‘Propeltidium color’: [0] no color, [1] color halfway down, [2] color down sides. 2 ‘Num- ber of ctenidia’: [0] zero, [1] two, [2] four. 3 ‘Ctenidia shape’: [0] NA, [1] flat, [2] peg, [3J thin. 4 ‘Ctenidia length’: [0] NA, [1] short, [2] medium, [3] long. 5 ‘Presence/absence palpal papillae’: [0] present, [1] absent. 6 ‘Pal- pal coloration’: [0] pale or dusky, [1] dark tib- ia and metatarsus, [2] dark tarsus, metatarsus, tibia, and femur. 7 ‘PIT’: [0] separate, [1] in notch, [2] NA. 8 ‘FF crimped’: [0] no crimp, [1] crimped. 9 ‘Cleft AT: [0] no, [1] yes, [2] NA. 10 ‘Size AT’: [0] small, [1] large, [2] NA. 11 ‘GO shape’: [0] smooth, [1] undulate, [2] hooked. 12 ‘MST’: [0] medium, [1] tiny, [2] absent. We used PAUP version 4.0 beta (Swofford 2002) to analyze the data. All characters were Linordered and given equal weight. We used the branch and bound (b and b) technique to search for the most parsimonious trees result- ing in one tree (Fig. 61) with a length of 38, a consistency index of 0.63 and a retention index of 0.68. Characters were then reweight- ed by the maximum value of the consistency indices resulting in the same tree. However, the length was reduced to 24, the consistency index was 0.68 and the retention index was 0.73. When the characters were reweighted, three characters (2, 5 & 8) had a weight of 1; six characters (1, 4, 7, 9, 10 and 11) had a weight of 0.67; two characters (3 & 12) had a weight of 0.5; and character 6 had a weight of 0.4. Although our cladistic analysis resulted in one best tree, the relatively low consistency index and the relatively few characters deemed useful for the cladistic analysis points to the need for exploring the use of molecular markers in future phylogenetic analyses of so- lifugids. DISCUSSION Characters previously utilized by Muma, i.e. coloration, number and shape of ctenidia, number of palpal papillae, length (depth) ver- sus width of fondal notch were valuable char- acters for differentiating species although sta- tistical analysis of the FL/FW ratio character demonstrates that in several species there is more variability than Muma indicates. In ad- dition the shape of the anterior tooth on the movable finger of the males, the presence and size of the mesal tooth and the position of the posterior intermediate tooth of the movable finger relative to the movable finger on both males and females was important. Some spe- cies also had a consistent cleft underneath the anterior tooth of the MF caused either by a depression of the dorsal edge of the FF or a slight ridge that formed anterior to the AT of the FF. Shape of male fixed finger from an ectal view was also important with some dis- playing an upward undulation or “crimp” and others a more normally rounded appearance. Figures 27-35 demonstrate variability and are presented for comparison purposes. In those populations in which sufficient specimens were available for examination there was some variation in the shape and number of ctenidia but useful general criteria could be established. Some members of the same species had ctenidia that were bifed while the rest were pointed or blunt, probably because of wear. Infrequently some ctenidia were bent (see Figs. 36-43 for comparison). Some female groups displayed ctenidia but in all cases they were thin and short (Fig. 44). We did not find them to be diagnostic. We examined penultimate specimens of E. similis from Sevilleta LTER for ctenidia but found none. In those species in which palpal papillae are part of the diagnosis there was some var- BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 309 pallipes actenidia hodai scaber dams ctenidieilus ascopulatiis zinni socal corpink mormonus icenoglei similis Figure 61. — Cladogram (using Paup 4.0) of the Eremobates species group with a length of 24, a consistency index of 0.68 and a retention index of 0.73, after character reweighting (see text for details). Eremobates pallipes was used as the outgroup. 310 THE JOURNAL OF ARACHNOLOGY Table 1 . — Character matrix for species of the scaber group of the genus Eremobates with E. pallipes (Say 1823) as the outgroup. See text for description of characters. 1 2 3 4 5 6 7 8 9 10 11 12 E. pallipes 0 0 0 0 0 0 0 0 0 0 0 0 E. scaber 2 1 1 1 0 0 0 1 0 0 1 2 E. hodai 1 1 1 1 0 0 0 1 0 0 1 1 E. clams 2 1 2 1 0 0 0 1 1 0 1 2 E. ctenidiellus 2 1 3 2 0 0 1 1 0 0 0 1 E. ascopulatus 2 1 2 2 0 1 0 1 0 0 0 2 E. zinni 1 2 1 1 0 1 0 0 1 0 2 2 E. socal 2 2 1 1 0 1 0 0 1 0 2 1 E. icenoglei 1 2 2 1 1 2 2 0 2 2 2 0 E. morrnonus 1 2 3 3 0 1 1 0 0 1 2 1 E. corpink 2 1 3 1 0 1 0 0 1 0 2 1 E. actenidia 1 0 0 0 0 1 0 0 0 1 1 1 E. similis 1 2 3 1 1 2 1 0 0 1 2 1 iation among the number of palpal papillae including an occasional complete absence. There is some evidence that papillae arise se- quentially after the last molt. Propeltidium coloration was also helpful in identifying several species. Many species had light brownish violet mottling on the anterior edge but several also had distinctive color pat- terns (see Figs. 45-51). The use of coloration in solifugids captured in pitfall traps and pre- served in alcohol should be approached with caution although this trait is used in this study. Material from Hanford, Washington contained both light and dark specimens in the same vial. Solifugids collected in wet pitfall traps from San Luis Valley, Colorado changed color over a two year period (Brookhart, pers. obs.). Female coloration was almost always the same as the males. Female genital opercula were quite species specific (Figs. 52-60). Male ratios of A/CP, FL/FW, FW/FFW proved to be statistically significant for some species. Using the Tukey-Kramer analysis of variance we found E. similis and E, morrnonus to be long legged species and E. hodai to be a short legged species. All others showed no significant statistical difference. Only E. icen- oglei proved to have a longer FL/FW ratio and E. similis and E. morrnonus were wider. The FW/FFW ratio identifies those species with wider or thinner fixed fingers. Those with sig- nificant differences were E. actenidia with a thin FF and E. clarus, E. icenoglei, and E. scaher with wider FF. All were significant at the 95% level of confidence. PL/PW and CL/ CW showed no significance. No statistically significant character was found among any of the female ratios tested. Species vary in the position of the posterior intermediate tooth relative to the primary tooth of the movable finger and the presence or absence of a cleft anterior to the anterior tooth of the movable finger. Brookhart & Muma (1981, 1987) and Muma & Brookhart (1988) demonstrated that solifugids are generally allopatric or in the case of the Eremobates palpisetulosus group sympatric for two species. Our study indicates that members of the scaber group are allopat- ric except E. ascopulatus and E. zinni. Our cladistic analysis seems to demonstrate a northern and a southern clade (compare the cladogram in Fig. 61 with the map in Fig. 62). Males in the northern clade have two ctenidia, a more pronounced ‘"crimp” in the fixed fin- ger of the male and females with either smooth or undulate medial margin of the gen- ital operculum. Their range is larger than the southern group which has species with more endemic status. Southern males have four cte- nidia, a more smoothly curved fixed finger, and females have a hooked process at the juncture of the medial margin and the wing of the genital operculum. The sympatry between E. zinni and E. ascopulatus may reflect a con- nection within the Lahontan Basin. We do not have enough data to indicate whether there is a temporal separation or some other process of niche partition. ACKNOWLEDGMENTS We would like to thank all who helped us in this project: Kristy Alexander and Adrian BROOKHART & CUSHING— SCABER GROUP OF THE GENUS EREMOBATES 311 130°W 50°N 120°W 110°W Figure 62. — Map of Eremobates scaber group species. 312 THE JOURNAL OF ARACHNOLOGY Kropp of the Denver Museum of Nature and Science for assistance with illustrations and maps; Jim Luhring, Cherry Creek High School mathematics department for running the statistics; and Irene Brookhart for help with the manuscript and in the field. Thanks also to Mark Harvey and anonymous review- ers who provided invaluable suggestions for improvements to the manuscript. LITERATURE CITED Allred, D. & M.H. Muma. 1971. Solpugids of the National Reactor Testing Station, Idaho. Great Basin Naturalist 3 1(3): 164-168. Banks, N. 1900. Synopses of North-American In- vertebrates, IX. The Scorpions, Solpugids, and Pedipalpi. American Naturalist 34:421-427. Brookhart, J.O. 1972. Solpugids (Arachnida) in Colorado. Southwestern Naturalist 17:31-34. Brookhart, J. & S. Brantly. 2000. Solpugids (Arach- nida) of the Sevilleta National Wildlife Refuge, New Mexico. Southwestern Naturalist 45:443- 449. Brookhart, J. & RE. Cushing. 2002. New species of Eremobatidae (Arachnida, Solifugae) from North America. Journal of Arachnology 30:84-97. Brookhart, J.O. & M.H. Muma. 1981. The pallipes species-group of Eremohates Banks (Solpugida: Arachnida). Florida Entomologist 64(2):283- 308. Brookhart, J.O. & M.H. Muma. 1987. Arenotlieriis, a new genus of Eremobatidae (Solpugida) in the United States. Pp. 1-18 and 4 plates, printed for the authors by Cherry Creek High School Print Shop, Englewood, Colorado. Hall, E.R. 1946. Mammals of Nevada. University of California Press, Berkeley, California. Harvey, M.S. 2002. Nomenclatural notes on Sol- fugae, Amblypygi, Uropygi and Araneae (Arach- nida). Records of the Western Australian Muse- um. 20:449-459. Knisley, C.B. & J.M. Hill. 2001. Biology and con- servation of the Coral Pink Sand Dunes tiger beetles {Cicindela linibcita albissima Rumpp). Western North American Naturalist 61:381-394. Koch, C.L. 1842. Systematische uebersicht iiber die familie der Galeoden. Archiv fiir Naturgeschi- chte. 8:350-356 Kraepelin, K. 1899. Zur Systematik der Solifugen. Mitteilungen aus dem Naturhistorischen Museum in Hamburg 16:197-259. Kraepelin, K. 1901. Palpigradi Und Solifugae. Das Tierreich 12:1-159. Muma, M.H. 1951. The arachnid order Solpugida in the United States. Bulletin of the American Museum of Natural History 97(2):35-141. Muma, M.H. 1962. The arachnid order Solpugida in the United States, Supplement 1, American Museum Novitates 2092:1-44. Muma, M.H. 1963. Solpugida of the Nevada Test Site. BYU Science Bulletin of Biology Series 3(2):1-15. Muma, M.H. 1970. A synoptic review of North American, Central American and West Indian Solpugida (Arthropoda: Arachnida): Arthropods of Florida and Neighboring Land Areas 5:1-62. Muma, M.H. 1985. A new possibly diagnostic char- acter for Solpugida (Arachnida). Novitates. Arthro- podae J-B Pub. Co., Crete, Nebraska 2(2): 1-5. Muma, M.H. 1987. New species and records of Sol- pugida (Arachnida) from Mexico, Central Amer- ica and the West Indies. Pp. 1-24, and 4 plates. Privately published for the author by Southwest Offset, Silver City, New Mexico. Muma, M.H. 1989. New species and records of Sol- pugida (Arachnida) from the United States. Pub- lished for the author by Douglas Print Shop, Douglas, Arizona. 50 pp., 8 plates. Muma, M.H. & J.O. Brookhart. 1988. The Eremo- bcites palpisetulosiis species-group (Solpugida: Eremobatidae) in the United States. Pp. 1-65, plates 1-20. Published for the authors by Cherry Creek High School Print Shop, Englewood, Col- orado. Punzo, E 1998. The Biology of Camel Spiders (Arachnida, Solifugae). Kluwer Academic Pub- lisher. Roewer, C.ER. 1934. Solifugae, Palpigradi, In Bronn, H. G. Klassen und Ordnungen des Tier- reiehs, Leipzig, vol. 5, div. 4, book 4, 723 pp. Simon, E. 1879. Essai d’une classification des Gal- eodes. Annales de la Societe Enetomologique de France, 5(9):93-154. Sokal R.R. & EJ. Rohlf 1981. Biometry. 2"^ Ed. W.H. Freeman and Company, New York, N.Y, USA. Swofford, D.L. 2002. PAUP*: Phylogenetic Anal- ysis Using Parsimony (and Other Methods), 4.0 Beta for Macintosh. Sinauer Associates. Trimble, S. 1989. The Sagebrush Ocean, University of Nevada Press, Reno. United States Environmental Protection Agency. 1986. Ecoregions of the Pacific Northwest. EPA/ 600/3-86/033. Vasquez, I. 1981. Contribucion al conocimiento de los solfugos de Mexico. (Arachnida: Solpugida). Tesis Profesionale. Universidad Nacional auto- noma Mexico. 82 pp. Manuscript received 24 Eebruary 2003, revised 21 August 2003. 2004. The Journal of Arachnology 32:313-323 ASSEMBLAGES OF SPIDERS ON MODELS OF SEMI-ARID SHRUBS Michael W* Heikkieen^ and James A. MacMahon: Department of Biology and the Ecology Center, Utah State University, 5305 Old Main Hill, Logan, Utah 84322- 5305, USA. E-mail: mheikkinen@boisestate.edu ABSTRACT. Many environmental factors influence the composition of animal assemblages. For spider assemblages, plant architecture is an important variable. Here we examine the effects of various plant architectural attributes by using models of shrubs in which we control branch orientation (horizontal or vertical) and height above the ground (0, 10, or 40 cm). Guild membership, based on hunting strategy (jumpers, pursuers, ambushers, or trappers), was used to characterize spider assemblages. Five replicates of the six treatments (two orientations by three heights) were randomly placed in a 60 m by 50 m grid among big sagebrush in a shrub-steppe habitat and sampled at 3 week intervals from July-October in 1997 and 1998. ANOVA was used to demonstrate that not only do single architectural variables influence the distribution of spiders but also the interaction of architectural variables influence spider distribution. Differences in the assemblages of spiders on the models were the result of architecture differences. Jumpers selected horizontal, 10 cm models and pursuers selected vertical, 0 cm models. Trappers were most abun- dant on horizontal, 0 cm models. Keywords: Community ecology, plant architecture, shrub- steppe The distributions of a wide variety of or- ganisms are influenced by structural charac- teristics of their physical environment (Mac- Arthur 1958; Wilson 1974; Rotenberry & Wiens 1980; James & Wame 1982; Landres & MacMahon 1983; Vander Wall & Mac- Mahon 1984; Southwood 1996). In particular, plant attributes correlate with animal species diversity (Schoener 1968; Pianka 1973; Law- ton 1986). Spiders have been the focus of many com- munity ecology studies because they are gen- eralist carnivores, many species live in the same habitat and they are easily collected (Wise 1993). Several habitat structures corre- late with spider abundance and diversity (Cole- bourn 1974; Gibson et al. 1992; Johnson 1995; Halley et al. 1996). Plant architecture was the specific subject of many studies (Fau- tin 1946; Chew 1961; Allred & Beck 1967; Allred 1969; Chaplin 1976; Guenarsson 1988; 1990, 1996; Janetos 1986; Ward & Lubin 1992; Wise 1993; Sundberg & Gunnarsson ’ Current address: Department of Curriculum, In- struction, and Foundational Studies, Boise State University, 1910 University Drive, Boise, Idaho, 83725-1745. 1994; Aiken & Coyle 2000; Ysnel & Canard 2000; Raizer & Amaral 2001). The effect of plant architecture on the dis- tribution of spiders on big sagebrush {Arte- misia tridentata) has been the focus of several studies. Architectural features, such as herb height and shrub size were associated with the distribution of spiders (Abraham 1983). Changes in the density of individual big sage- brush altered the composition of the spider community (Hatley & MacMahon 1980; Wing 1984). Robinson (1981) and Ehmann (1994c) used models to simulate big sagebrush den- sity, substrate diameter and horizontal and vertical orientation. Guilds, based on mode of feeding (Root 1967), are widely used as dependent variables in studies of spider assemblages (Chew 1961; MacMahon 1973; Uetz 1977; Moran & South- wood 1982; Hurd & Eisenberg 1990; Petters- son 1996; Mason et al. 1997), Guild analysis provides a way to examine the organization of spider communities on big sagebrush (Hatley & MacMahon 1980; Robinson 1981; Abra- ham 1983; Wing 1984; Ehmann 1994a). Pre- vious studies suggest that the guild composi- tion of spider assemblages on big sagebrush is predictable despite differences in species 313 314 THE JOURNAL OF ARACHNOLOGY composition (Abraham 1983; Ehmann 1994a). Although jumpers are ubiquitous and are found on the ground and on most vegetation, they dominate the spider assemblages found on big sagebrush. In the guild classification used by Ehmann, the spider community on big sagebrush was divided into four guilds (trap- pers, jumpers, ambushers and pursuers) using hunting strategy as a means of identifying guild membership. Trappers, including Ara- neidae, Dictynidae, Linyphiidae, Tetragnathi- dae and Theridiidae, construct webs to trap prey. Jumpers, Oxyopidae and Salticidae, ac- tively seek prey using their well developed sense of sight to pounce on prey from a dis- tance. Ambushers, Thomisidae, are sit-and- wait predators that wait for prey to move within striking distance. Pursuers, including Anyphaenidae, Clubionidae, Gnaphosidae, Lycosidae and Philodromidae, are active pred- ators that run down prey. In this study, the distribution of spiders by guild and a possible mechanism for the ob- served patterns were addressed using an ex- perimental approach in the well-studied spider assemblage on big sagebrush. Two shrub char- acteristics stand out as making them architec- turally different than the surrounding plants in shrub-steppe ecosystems. They are taller and they have more horizontal plant components. Shrub height has been associated with differ- ences in spider community composition (Hat- ley & MacMahon 1980; Abraham 1983; Greenstone 1984; Dobel et al. 1990; Gibson et al. 1992 ; Lubin et al. 1993; Ward & Lubin 1993; Aiken & Coyle 2000; McReynolds 2000). Robinson (1981) used simple models to demonstrate that vertical/horizontal orien- tation affected the distribution of certain spi- der species. In this study, artificial shrubs were used to manipulate height and orientation of pseudobranches to determine the roles these two variables play in the distribution of spi- ders affiliated with specific guilds. Data from a four-year census of spiders on big sagebrush in the same area as the experiment were used to interpret results in relation to the natural system. The specific purpose of this study was to measure the distribution of spiders affiliated with each of four guilds on models that sim- ulated two plant architectural variables: height and branch orientation. Spider abundance on each model served as the dependent variable in the analyses. METHODS Site. — This study was conducted at a 10 ha site 3.7 km east and 0.9 km north of the Hyde Park, Utah post office (NWl/4 SWl/4 sec 6, T 12 N, R 2 E, Salt Lake Meridian) at an elevation of 1755 m. The site was dominated by big sagebrush and grass with alfalfa fields on the east and west margins and steep can- yons on the north and south. The experiment took place on the south side of a farm road that divided the site. This area has a southwest aspect and 5% slope. The part of the site north of the road was used for a four-year census. This site is 4 km north of the area used by Hatley and MacMahon (1980), Robinson (1981), Abraham (1983), and Wing (1984) and 7.5 km northwest of Ehmann's site (1994a). 4-year census. — An 80 m^ grid was estab- lished and divided into quadrants to facilitate locating sampling points. The four corners, midpoints of each side and the center were permanently marked. In 1995, 20 shrubs were chosen for the census for each sampling day. After 1995 the number of shrubs sampled per day was increased to 24. Spiders were col- lected every 14-28 d from May until October in 1995-98. The number of sample days ranged from 7-13 totaling 42 for the four years. For each census day, five or six points within the grid were selected using a table of random numbers to identify the coordinates of the points. Sampling began 2 h after sunrise using the randomly selected coordinates. The four big sagebrush closest to the sampling point and meeting the established criterion (0.75-1.5 m high) were noted. The height cri- terion was established in order to sample sim- ilar sized big sagebrush because there is a pos- itive relation between shrub size and spider abundance (Abraham 1983). Spiders were collected from each shrub by using a beating sheet technique (Southwood 1978; Ehmann 1994a). Large spiders were captured by hand using vials and small spiders were captured with an aspirator. The beating sheet technique captures about 84% of the spi- ders on big sagebrush and the capture rate rep- resents an unbiased sample of species found on the shrubs (Ehmann 1994b). Ehmann also showed that their was no effect on the sub- HEIKKINEN & MACMAHON— ASSEMBLAGES OF SPIDERS 315 Figure 1. — Photograph of model in horizontal orientation. sequent sampling of the same shrub after a two-week interval. All spiders captured from a shrub were immediately placed in a vial with 70% ethanol for preservation. This pro- cess continued until the four shrubs around the first point were sampled. The second point was then located and the process repeated un- til all 24 shrubs were sampled. Spiders were sorted by species and identi- fied using the reference collection of Cache Valley spiders at Utah State University (Eh- mann 1994b). Data were recorded as total spe- cies abundance and guild abundance per day. Guild assignments (jumpers, pursuers, am- bushers, and trappers) followed the proce- dures used by Ehmann (1994a). Specimens were deposited in the Utah State University Entomology Collection. A pre-experiment census was also conducted, in the same man- ner, for one season (1997) in the area where the experimental models were placed. The Jaccard index, a coefficient of similar- ity that gives an indication of the degree to which species composition overlaps between two locations (Southwood 1978), was used to compare assemblages of spiders in the exper- iment with the natural distribution found on big sagebrush. Experiment with models. — The models for the experiment were constructed from wood and sisal twine and in some cases alu- minum conduit (Eigs. 1 & 2). A center post was the “trunk.” A “whorl” of eight dowels “branched” perpendicularly from each end. A wooden slat was used to keep the ends of the Figure 2. — Photograph of model in vertical ori- entation. branches 40 cm apart. Sisal twine wrapped around the dowels was used to represent smaller branches. The models were cylinders with 63 cm diameters, 40 cm heights and vol- umes of 124,690 cm^ (Heikkinen 2001). Shrub volumes measured in previous studies were 24,991-598,796 cm^ (Hatley & MacMahon 1980) and 120,000-311,000 cm^ (Wing 1984). Two orientations were tested (vertical and horizontal) by turning the model 90° on its longitudinal axis. Three height positions were used for each of the vertical/horizontal orientations. One treatment was placed on the ground. In the second and third treatments aluminum conduit was used to raise the model 10 cm and 40 cm off the surface. The conduit was driven into the ground and then inserted in a hole drilled into the center post. Five replicates of each treatment were established on a 10 m by 10 m grid. Treatments were assigned to each point on the grid by random draw. Once a treatment was assigned to a point, it remained at that point for the year, but a second random assignment took place for the second year. Where sample points coincided with a shrub, the model was placed as close to the sample point as possible without touching a shrub. Data were collected in 1997 and 1998 be- ginning in the last week of July and ending in mid-October. Spiders were collected from the models every 3 weeks using the beating sheet technique, and preserved in 70% ethanol. Specimens were sorted and identified in the laboratory and assigned to one of the four 316 THE JOURNAL OF ARACHNOLOGY guilds using the same procedures described above. Data were analyzed using ANOVA (SAS 1982) where species the independent variable was an individual spider and the dependent variables were the spider’s guild, the orienta- tion of the model on which it was found, and the height of the model. All two-way and three-way interactions were tested. RESULTS Censuses of spiders on big sagebrush at Hyde Park. — A baseline for the interpretation of these experiments was established from censuses of spiders found on big sagebrush from 1995-1998 (Table 1). The jumper guild was most abundant each year containing 62.0% of the total individuals (range = 57.5- 66.0%). Four jumpers were among the ten most abundant species across all guilds. Pe~ legrina aeneola Curtis 1892, a jumper, was most abundant accounting for 26.9% of the total. Other abundant jumpers were Sassacus papenhoei Peckham & Peckham 1895, Oxy- opes scalaris Hentz 1894 and Phidippus john- soni Peckham & Peckham 1883. The trapper guild was second most abundant with 21.1% of the individuals collected and it had three of the ten most abundant species: Theridion pe- traem L. Koch 1872, T. neomexicanum and Metapeira foxi Gertsch & Ivie 1936. Pursuers was the third most abundant guild with 12.9% of the total. The most abundant pursuers were Philodromus histrio Latreille 1819 and Tibel- lus oblongus Walckanaer 1802. Ambushers was the least abundant guild with only 2.9% of the spiders collected. One ambusher, Xys- ticus gulosus Keyserling 1880, was the tenth most abundant species. Results of the experiment with models. — The Jaccard index yielded a similarity value of 0.55 between the Hyde Park census and the experiment. See Table 1 for a list of the spe- cies found in the census and the experiment. Our principle hypothesis was that jumpers would be most abundant on shrub-like mod- els, i.e., 40 cm models placed in a horizontal orientation. Since this was an experimental manipulation of two shrub variables which may effect the distribution patterns of all spi- der guilds, the significance of all variables (height, orientation and guild affiliation) and higher-order interactions were identified using ANOVA (Table 2). Two variables were significant by them- selves. Over half of all spiders collected were jumpers and about half of all spiders were on the 0 cm models (Table 3). Although the assemblages of spiders based on guild membership from the censuses and the experiment were similar (Fig. 3), there were differences in the species composition of the assemblages. Jumpers was the most abun- dant guild in both cases (65.8% and 51.6%), however, Pelligrina aeneola was most abun- dant in the census, but accounted for only I. 4% of all spiders on the experimental mod- els. Although pursuers were third most abun- dant in both treatments, they accounted for I I. 0% of the spiders in the census, but 19.4% of the spiders collected from the experimental models. Philodromus histrio was the most abundant pursuer in the census (4.8%), but was only 0.2% of the spiders collected from the experimental models. Tibellus oblongus was more abundant on the experimental mod- els: 17.9% vs. 4.2%. The significant variables identified by the ANOVA only indicate that spiders are react- ing differentially to architectural variables. Two-way interactions were examined to elu- cidate differences in spider distribution by guild based on differences in height or ori- entation. Even though orientation was not sig- nificant alone, height by orientation was also included because the complete model was an- alyzed. All two-way interactions were signif- icant. The guild by height interaction was signif- icant because 86% of the pursuers and 44% of the trappers were on the 0 cm models. Ori- entation by itself was not a significant vari- able, about half were on models of each ori- entation; the interaction of orientation with guild and height was significant. Jumpers were more abundant on horizontal models (58%) and pursuers were more abundant on vertical models (81%). The height by orien- tation interaction was significant because 61% of the spiders on the 10 cm models were on horizontal ones and 57% of spiders on the 0 cm models were on vertical ones. The three-way interaction was the analysis used to test our principle hypothesis that spi- ders belonging to specific guilds would be more likely to be found on specific models. For example, jumpers would be most abun- dant on tall, horizontal models. The three-way HEIKKINEN & MACMAHON— ASSEMBLAGES OF SPIDERS 317 Table 1 . — Abundance of spider species found at Hyde Park and on the experimental models. Species Census Experiment 1995 1996 1997 1998 1997 1998 Sassacus papenhoei 267 707 143 196 19 31 Pelegrina aeneola 251 966 433 717 4 2 Phidippus johnsoni 49 121 42 58 14 20 Evarcha hoyi 15 10 39 63 1 1 Habronattus hirsutus 13 15 1 0 0 0 Tutelina similis 2 12 4 19 2 4 Pellenes hirsutus 3 27 5 10 0 4 Synagaies idahoansis 0 12 1 1 0 1 Talavera sp. 0 2 2 6 0 1 Oxyopes scalaris 221 628 81 306 44 85 Salticidae 0 10 4 0 0 2 Jumpers 821 2510 755 1376 74 147 Philodromus histrio 68 235 53 181 0 1 PMlodromus rufus 0 24 5 4 1 0 Philodromus sp. 0 1 1 1 0 0 Tibellus oblongus 71 188 58 95 33 44 Ebo evanses 0 0 0 1 0 0 Thanatus formicinus 2 0 0 0 0 0 Cheiracanthium inclusum 5 74 23 23 2 2 Anypkaena pacifica 1 3 1 0 0 0 Zelotes subterraneus 3 0 0 1 0 0 Gnaphosidae 5 1 1 2 0 0 Clubionidae 0 1 1 2 0 0 Unknown pursuer 0 1 1 0 0 0 Pursuers 155 528 144 310 36 47 Xysticus guiosus 21 116 12 28 5 7 Xysticus cunctator 0 2 0 0 0 0 Xysticus montanensis 2 1 0 0 0 0 Xysticus sp. 2 0 0 1 0 0 Misumenops lepidus 26 6 7 13 1 2 Misumenoides sp. 0 0 1 0 0 0 Coriarchne utahensis 6 2 2 4 0 1 Ambushers 57 127 22 46 6 10 Theridion petraeum 124 305 142 229 9 42 Theridion neomexicanum 36 61 104 57 11 1 Theridion differens 0 0 1 4 0 0 Theridion sp. 25 8 5 10 2 1 Euryopsis scriptipes 20 58 30 57 2 8 Enoplognatha ovata 0 3 1 3 0 1 Diponea tibialis 0 38 20 12 2 0 Diponea nigra 0 11 9 2 0 0 Dictyna completa 10 1 0 12 0 0 Dictyna idahoana 18 33 9 6 1 1 Metepeira foxi 27 85 50 73 12 11 Erigone dentosa 27 21 4 40 0 0 Spirembolus mundus 3 2 2 1 0 0 Frontinella communis 2 3 1 3 1 0 Araneus gemma 1 6 2 5 0 0 Araneus displicatus 1 4 1 4 0 1 Aculepeira verae 0 5 1 3 0 0 Aranidae 0 1 0 0 0 0 Linyphidae 0 0 0 3 0 0 Unknown trapper 0 4 6 3 2 1 Trappers 295 647 388 527 42 67 Unknown 93 4 3 1 0 0 Total 1421 3806 1312 2263 158 271 318 THE JOURNAL OF ARACHNOLOGY Table 2. — Results of the ANOVA of spider abundance on the experimental models. Source df SS MS F P Replicates Guild Height Orientation 1 Guild X Height 6 Guild X Orientation 3 Height X Orientation 2 Guild X Height X Orientation 6 En-or 816 1.210 0.303 0.45 0.7733 101.927 33.976 50.43 0.0001 25.474 12.737 18.91 0.0001 0.001 0.001 0.00 0.9666 22.526 3.754 5.57 0.0001 17.899 5.966 8.86 0.0001 4.617 2.308 3.43 0.0330 23.269 3.878 5.76 0.0001 547.019 0.674 interaction was significant, and jumpers were most abundant on the 10 cm horizontal mod- els (Fig. 4). This meant that there were unique combinations of height and orientation which had greater abundances of spiders from par- ticular guilds. The most obvious interaction was that 70% of all pursuers were on the 0 cm vertical models. Jumpers were about equally abundant on all model types, but the effect of the interaction is evident when the abundances for the 10 cm models are com- pared. Jumpers were most abundant on the 10 cm horizontal models and least abundant on the 10 cm vertical models. Twenty-nine per- cent of all trappers were on 0 cm horizontal models. DISCUSSION The spider community found on big sage- brush has a characteristic distribution of spi- der guilds that is dominated by jumpers (Eh- mann 1994a; Abraham 1983). Previous studies suggested that plant architecture influ- enced guild abundance of spiders on big sage- brush (Ehmann 1994a; Wing 1984; Robinson 1981; Hatley & MacMahon 1980). In this study, the effects of two architectural variables on the distribution of spider guilds was tested using models that simulated the volume, branch texture and branch diameter of big sagebrush, while keeping the structural details simple enough to measure the two treatment variables (height above ground and branch orientation) and control for other variables. The models were placed among big sage- brush, so they were in the right habitat, and since spiders readily disperse (Dean & Ster- ling 1985; Bishop & Riechert 1990; Ehmann 1994b; Foelix 1996), they were in a habitat which contained a pool of potential colonists. Previous studies that examined habitat com- plexity, looked at the effect of single variables on the distribution of spiders. In this study, it was possible to test for the significance of the height by orientation interaction on the distri- bution of spider guilds. The significant three- way interaction indicated that branch orienta- tion and height had a differential effect on guild abundance. Table 3. — Total guild abundance for each model type over the two year (1997-1998) experiment using models to simulate shrub architecture. Numbers are the totals for seven sampling periods. Percentage of total abundance is in parentheses. 0 cm, 10 cm, and 40 cm = three height treatments. H = horizontal orientation and V = vertical. 0 cm 10 cm 40 cm Guild Guild H V H V H V totals Jumper 44 (10.2) 41 (9.5) 54 (12.5) 22 (5.1) 31 (7.2) 29 (6.7) 221 (51.3) Pursuer 13 (3.0) 58 (13.5) 1 (0.2) 7 (1.6) 2 (0.5) 2 (0.5) 83 (19.3) Ambusher 2 (0.5) 5 (1.2) 3 (0.7) 2 (0.5) 5 (1.2) 1 (0.2) 18 (4.2) Trapper 32 (7.4) 16 (3.7) 15 (3.5) 16 (3.7) 1 1 (2.6) 19 (4.4) 109 (25.3) Model totals Height totals 91 (21.1) 211 (49.0) 120 (27.8) 73 (16.7) 120 (27.8) 47 (10.9) 49 (11.4) 100 (23.2) 51 (11.8) HEIKKINEN & MACMAHON— ASSEMBLAGES OF SPIDERS 319 Guild Distribution 70 60 Jumpers Pursuers Ambushers Trappers Guild Figure 3. — Guild distribution of spiders from the Hyde Park census and from the experiment. Abundance ■ Jumper ID Trapper □ Pursuer B Ambusher Figure 4. — Abundance of spiders by guild on models at two orientations (V = vertical and H = Horizontal) and three heights (0, 10, and 40 cm). 320 THE JOURNAL OF ARACHNOLOGY Seventy percent of all pursuers were found on the 0 cm, vertical models (Fig. 4). This model probably simulated grasses, and the most abundant pursuer in the experiment was T. oblongus, a grass specialist according to Roberts (1995), that is usually described as a sit-and-wait predator more like an ambusher than the other members of its guild (Gertsch 1979). The most common pursuer on big sagebrush at Hyde Park was P. histrio, but only one was collected from the models in the experiment. Salticids and oxyopids are often more abun- dant on one vegetation type than another (Cut- ler et al. 1977; Abraham 1983). Jumpers are dominant on big sagebrush as they were on the horizontal models in the experiment. Jumpers were found in high numbers on all of the models, and there appears to be no dif- ferences between orientations on the 0 cm and 40 cm models. However, 71% of the jumpers on the 40 cm models were on those with hor- izontal orientations (Fig. 4). Sagebrush has significant horizontal components, so the hor- izontal nature of sagebrush may be one of the architectural variables to which jumpers are responding when they choose to remain on a shrub. The two-way interaction showed jump- ers preferred horizontal models. In this exper- iment, jumpers were 51.4% of all spiders, which was within the range of jumpers found on big sagebrush in Hyde Park, but the most common jumper, P. aeneola, was virtually ab- sent in the experiment. The decrease in num- bers of P. aeneola was offset by higher num- bers of O. scalahs. We do not know why there were species replacements. The important point is that there are redundant species and the guild distribution remained similar. Salticids are among the most neurally so- phisticated spiders (Forster 1982). The com- bination of a keen sense of sight, the ability to track prey even when the line-of-sight is interrupted and their unique jumping ability make them particularly adept at hunting in the structurally complex habitat found inside shrubs (Land 1969; Enders 1975; Jackson 1986; Jackson & Tarsitano 1993). Jumpers may be using horizontality as a mechanism for recognizing that they are in a shrub, selecting habitat based on environmen- tal cues (Orians & Wittenberg 1991). To un- derstand these relationships additional studies are needed of the mechanisms responsible for these responses (Rypstra et al. 1999) and the life history and foraging behavior of these species (Neuvonen 1999). The three-way interaction for trappers dem- onstrates a more complicated height by ori- entation interaction. Forty-four percent of all trappers were on the 0 cm models. Of those, two-thirds were on the horizontal models. This result agrees with Robinson’s (1981) finding. A interesting feature of this three-way interaction is that an opposite result was found for the models placed 40-cm above the ground. Twenty-eight percent of all trappers were on these models, but, in this case, 63% of them were on the vertical models. The distribution of orb-web weaving trap- pers and cob-web weaving trappers (using Abraham’s (1983) designations) also differed between these two treatments, with a higher proportion of orb-weavers on the taller verti- cal models and a higher proportion of cob- weavers on the horizontal ground models. Per- haps placing all spiders that use webs as snares in the same guild is too simplistic. Uetz et al. (1999) recently divided trappers into more than one guild. Ambushers play a minor role on big sage- brush, as was true in the experiment. Am- bushers are sit-and-wait predators. Many sit in the flowers of plants waiting to ambush pol- linators. The small flowers of big sagebrush do not provide good sites from which to am- bush prey. The majority of ambushers on big sagebrush are probably using crevices in the bark as retreats. The significant three-way interaction dem- onstrates that spider decision-making involves a complex integration of environmental cues. The models were purposely simplified so the two variables of interest could be experimen- tally manipulated. The simplification had the effect of eliminating other variables, which are characteristic of big sagebrush, that spi- ders may also use as cues in the decision-mak- ing process. Some of these variables are bark texture, leaf structure, color, phytochemicals, and structural complexity. The result from this experiment that two of the most prominent members of the Hyde Park big sagebrush spi- der community, P. aeneola and P. histrio, were virtually absent on the experimental models in both years, indicates that the deci- sion-making process probably involves more than two variables. HEIKKINEN & MACMAHON— ASSEMBLAGES OF SPIDERS 321 The experimental nature of this study made it possible to establish the cause and effect relationship among the architectural variables and the distribution of spider guilds„ Spiders use architectural cues as part of the decision- making process to establish residency on shrubs or to make an attempt at colonization elsewhere, LITERATURE CITED Abraham, BJ. 1983. Spatial and temporal patterns in a sagebrush steppe community (Arachnida, Araneae). Journal of Arachnology 11:31-50. Aiken M. & EA. Coyle. 2000. Habitat distribution, life history and behavior of Tetragnatha spider species in the Great Smoky Mountains National Park. 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The Journal of Arachnology 32:324-331 GROUP SIZE DOES NOT INFLUENCE GROWTH IN THE THERAPHOSID SPIDER HYSTEROCRATES GIGAS (ARANEAE, THERAPHOSID AE, EUMENOPHORINAE) Melissa M. Varrecchia, Vanessa A. Gorley and Samuel D. Marshall': J. H. Barrow Field Station, Department of Biology, Hiram College, Hiram, OH 44234. ABSTRACT. Spiderlings of the theraphosid spider Hysterocrates gigcis were reared for 12 weeks with a superabundance of prey solitarily and in groups of two and four to examine the influence of rearing group size on growth. This taxon was selected because observations made on captive populations indicate that Hysterocrates spp. tarantulas have an unusually high level of mutual tolerance and captive juveniles have been observed to feed cooperatively on large prey until several months old. Cannibalism was only observed in one instance, in a group of four. There was no significant effect of rearing group size on increase in body mass. There was a tendency for a greater asymmetry in final weight in dyads than in tetrads. No difference was found in the amount of time spent feeding by individuals between the different group sizes. Hence, benefits of group living in Hysterocrates gigas spiderlings were not evident in this study. Keywords: Tarantula, sociality, Mygalomorphae Sociality in arachnids is a relatively rare phenomenon. Of approximately 36,000 de- scribed spider species, it is thought that only 35 are social; however, sociality has been demonstrated in at least 18 families of Ara- neae (Curtis & Can'el 1999). There are com- peting classification schemes for spider soci- ality (Aviles 1997), and the spectrum of spider sociality ranges from mutual tolerance to ac- tive cooperation in prey capture and brood care. One of the most significant thresholds along this continuum is the appearance of co- habitation by ecologically-competent juvenile spiders (i.e., those that could survive solitari- ly). These types of prolonged sibling aggre- gations are thought to represent an evolution- ary step in the direction of quasisocial behavior in which sexually mature spiders ex- hibit cooperative behavior (Aviles 1997). Sociality in spiders has presumably devel- oped because of the benefits that come with direct cooperation and sharing the costs of silk production. Benefits of group living could in- clude an increase in the amount and/or size of prey captured, shared construction costs of the web or increased predator avoidance (Aviles 1997; Uetz & Hieber 1997). Potential costs ‘ Current address: Dept, of Biological Science, Uni- versity of Cincinnati, Cincinnati, OH 45220 include direct competition for prey, increased predation or increased egg sac parasitism (Uetz & Hieber 1997). Sociality has been widely studied in the ar- aneomorph spiders, but has remained relative- ly unexamined in the mygalomorphs (Aviles 1997). Jantschke and Nentwig (2001) ob- served females of the subsocial diplurid spider Ischnothele caudata Ausserer 1875 caring for spiderlings by catching and sharing prey. The study by Darchen (1967) on the ischnocoline tarantula Heterothele darcheni (Benoit 1966) is the only documented case of sociality in theraphosids of which we are aware. Darchen found that these spiders display no aggression towards others in a group web, though they do not cooperate in hunting (Darchen 1967). Our study involved the theraphosid spider Hysterocrates gigas Pocock 1897. This West African tarantula lives in deep burrows in the rain forests and grasslands of Nigeria, Came- roon and the Congo (Smith 1990; Marshall 1996). Hysterocrates gigas buiTows have been found in a wide variety of locations: at the base of trees, beneath rotted logs, in termite mounds, on roadside embankments, on the pe- riphery of village compounds and on flat ground amongst palm groves, heavy grassland brush or dense tropical wet forests (Smith 1990; R. West pers. comm.). 324 VARRECCHIA ET AL.— GROUP SIZE AND GROWTH IN H. GIGAS 325 In this study we observed social interac- tions of H. gigas spiderlings in captivity. In captivity, we have observed this species shar- ing prey as juveniles up to several months of age. Although there have been no studies of H. gigas phenology in the wild, captive indi- viduals may reach maturity in 18-24 months (S. Marshall pers. obs.). Observations in the field in Cameroon show that well-grown young will cohabit in the maternal bun'ow with the mother (R. West, pers. comm.). In an 1 1 week study conducted by Reichling & Gutzke (unpub. data), spiderlings of the close- ly-related H. crassipes Pocock 1897 swarmed together on a prey item subdued by the moth- er, which was then completely devoured. Reichling and Gutzke’s observations demon- strate that sociality in Hysterocrates siblings can extend well beyond the first instar. In our study, sociality was observed for 12 weeks. We examined how group size (singles, dyads and tetrads) affected weight gain and feeding behavior in juvenile H. gigas. Because we had observed extended cohabitation of juveniles in captivity, as well as a unique group feeding behavior (i.e., cluster feeding) we predicted that H. gigas spiderlings reared communally would grow faster than those reared in isola- tion. METHODS Test subjects were obtained from two dif- ferent clutches of spiderlings produced by two wild-caught females collected in Cameroon and purchased from a commercial dealer (vouchers will be deposited at the American Museum of Natural History). The spiderlings remained in their communal sibling groups until we divided them into treatment groups. The treatment group sizes consisted of sib- lings placed as singletons, dyads or tetrads. The first clutch yielded seven replicates per group size and the second clutch yielded two replicates per group size. To differentiate in- dividuals within groups, each spiderling (sin- gletons included) was paint-marked on the dorsal side of the abdomen using Testors® enamel paint. At the beginning of the study, spiderlings from the two clutches differed in mass (Mean mg ± 1 SD: Clutch 1; 42.8 ± 14.2, n = 49, Clutch 2; 53.6 ± 18.7, n = 14). Because spiderlings within clutches were ran- domly assigned to treatment groups, this dif- ference in starting mass was not associated with treatment group size (ANOVA on spi- derling starting mass: Clutch, F, 57 = 8.82, P = 0.004; Treatment Group Size, F2 57 = 1 .74, P = 0.185). Group Size and Weight Gain. — Spiders were housed in translucent plastic 122 ml con- diment containers in a 14 L: 10 D cycle. The room was kept at an average temperature of 26.6 °C (range: 23 — 31 °C) and average hu- midity of 44.1 % RH (range: 30-63%RH). Substratum was not provided in the rearing container in order to facilitate observation and collection of prey remains. The spiderlings were given approximately 2 ml of distilled water each week in the bottom of the contain- er. The spiderlings were fed once a week. Prey consisted of pre-killed (by freezing) crickets ranging in weight from 100-450 mg. All treatments received the same size class of cricket at each feeding. This cricket size in- sured that food would always be in overabun- dance, eliminating food competition between spiderlings. Pre-killed prey was offered so that very large prey items could be used, items too large for the spiderlings to subdue. In a pilot study it was determined that spiderlings would feed readily on pre-killed prey. Superabun- dance of prey was verified by the presence of uneaten prey remains, which were collected 24 hours after feeding. Each spiderling was weighed to the nearest 0.1 mg in a tared plastic vial on an electronic balance before feeding and approximately 24 hours after feeding. The weights of the spi- derlings were recorded for a period of 12 con- secutive weeks. The average weekly weight gain was calculated within groups as was the coefficient of variation in weight gain for the last 4 weeks. Coefficients of variation are used to standardize variation in order to compare standard deviations of different sample sizes. A repeated measures ANOVA was used to as- sess the effect of group size on growth rates over the twelve weeks. Behavioral Mechanisms. — Details of feeding behavior were observed from week 9 until the termination of the study at week 12. Behavioral observations were conducted to in- vestigate any differences in time spent feeding between the spiders of the different sized treatment groups, any agonistic behaviors and occurrences of cluster feeding. Scan sampling was utilized to record the behavior of all spi- 326 THE JOURNAL OF ARACHNOLOGY Week of Study Figure 1. — Average weekly weight of Hysterocrates gigas spiderlings from two different maternal clutches in three treatment group sizes (singletons, dyads and tetrads). derlings during feeding. The frozen prey was introduced into each rearing container in the afternoon (between 1400-1600 hours). Scan samples were taken once every hour from the introduction of prey until midnight. Spider- lings were observed under red light after dark, unless differentiating the color marks of the spiderlings was difhcult, whereupon dim white illumination was used until the scan was complete. The frequency of feeding behavior was compared across the different group sizes. To compare feeding behavior and weight gain, the proportion of hourly intervals during which feeding was observed was arc sine square root transformed to normalize the data. The transformed proportion of hourly inter- vals during which feeding was observed cor- related with percent weight gain using a one- tailed Pearson’s r. A repeated measures ANOVA was used to determine differences in amount of time spent feeding among treat- ment groups. For all statistics an alpha level of 0.05 was used. RESULTS Group Size and Weight Gain. — For 12 weeks all groups of spiders gained weight at about the same rate. We found no significant effect of rearing group size on mass (repeated- measures ANOVA ; F, 2 = 0.08, P = 0.925). The average weekly weights showed no dis- tinguishable trend in any one group (Fig. 1). The variability in weight gain as expressed by the coefficient of variation for the last four weeks also showed no discernable trend for any one group (Fig. 2). Behavioral Mechanisms. — We observed the eight replicates of singletons, eight repli- cates of dyads and the five replicates of tetrads that remained at the end of the eighth week (unexplained mortality led to the loss of rep- licates during the course of the study). We re- corded these behaviors: Investigating, Feed- ing, Antagonizing, Grooming and Cluster Feeding. Investigating was defined as the spi- derling approaching the prey, contacting the cricket with the spiderling’s front legs, but not commencing in consuming the prey item. Feeding was defined as the spiderling con- tacting the cricket with its mouthparts. Antag- onizing was defined as: 1) chase, one spider- ling chasing another spiderling around the container, 2) kick, kicking another spiderling VARRECCHIA ET AL — GROUP SIZE AND GROWTH IN H. GIGAS 327 (0 > o E o o o 180 160 140 120 100 80 60 40 20 0 □ Singletons ■ Dyads □ Tetrads Week 9 Week 10 Week 11 Week 12 Week of Study Figure 2. — Variability in weight gain in Hysterocrates gigas spiderlings for the final four weeks of the study. No difference in the coefficient of variation of percent weight gain was found between groups. away from the cricket with its hind legs, or 3) take food, pulling the cricket away from the other spiderlings. Grooming was defined as a spiderling rubbing its legs together, over its abdomen, or over its cephalothorax. Cluster Feeding was defined as multiple spiderlings feeding on the same prey item at the same time, legs intertwined and no movement ob- served. The most common feeding behavior we ob- served among tetrads was spiderlings feeding individually (Table 1). The next most common behavior was two spiders feeding at the same time, but not in contact with each other. Clus- ter feeding was only observed on 7 occasions (6.9% of observations). All occurrences of cluster feeding were observed in tetrads. Tet- rads cluster-fed in groups of two, three or four. We found a correlation between individual weight gain and the percent of observations Table 1. — Feeding group sizes for Hysterocrates gigas spiderlings in tetrads. Tetrads were observed to feed in different sized groups. Number of occurrences lists number of hourly intervals. Feeding groups in tetrads Number of groups in which incident was observed Number of occurrences Percentage of occurrences One spider feeding 5 79 77.5 Two spiders feeding separately 5 15 14.7 Three spiders cluster feeding 3 3 2.9 Four spiders cluster feeding 3 3 2.9 Two spiders cluster feeding 1 1 1.0 Four spiders feeding separately 1 1 1.0 328 THE JOURNAL OF ARACHNOLOGY 0.35 ® Singletons H Dyads □ Tetrads Week 11 Week 12 Weeks Figure 3. — Proportion of hourly intervals during which Hyste roc rates gigas spiderlings were observed feeding by group size per week (mean +1 standard error). during which an individual was observed feeding (Pearson’s r: singletons, n = 32, r = 0.488, P = 0.002; dyads, n = 64, r = 0.477, P < 0.001; tetrads, n = 80, r = 0.176, P = 0.059). Using the Bonfen'oni adjustment for multiple correlations, an alpha level of 0.017 was set. For both the singles and the dyads, proportion of time spent feeding was signifi- cantly coiTelated with percent weight gain. No significant correlation was found in the tet- rads. We found that the proportion of time ob- served feeding was significantly different over the four weeks for the three treatment groups (repeated-measures ANOVA: F, 3 = 3.077, P = 0.035), showing that individual spiders dif- fered each week in the amount of time they were observed feeding. However, the be- tween-groups comparison indicated that the three treatment groups did not significantly differ from each other in proportion of time observed feeding (F, (, = 0.754, P = 0.609) (Fig. 3). A curious difference in mass was noted be- tween individuals in dyads. Coefficients of variation of the spiderlings’ final body weights at 1 2 weeks of age were used to com- pare the differences in spider size between dy- ads and tetrads. However, no significant dif- ference was found between coefficients of variation of body mass of dyads and tetrads at week 12 {t = 0.781, df = 9, P = 0.46). We did find a significant positive correlation of coefficient of variation of final body weights and the number of aggressive incidents ob- served (Speaman’s rho = 0.492, P = 0.044, n =13). DISCUSSION Over the 12 weeks of the experiment, spi- ders in each treatment group exhibited similar mass. We had anticipated a positive effect of group rearing based on the putatively adaptive cluster feeding behavior we observed because this feeding configuration appears to facilitate group feeding. So, why didn’t we find a growth-related benefit associated with com- munal rearing conditions? It may be that the superabundance of food provided by design allowed all spiderlings to feed to satiation. VARRECCHIA ET AL.— GROUP SIZE AND GROWTH IN H. GIGAS 329 Krafft et aL (1986) demonstrated that conspe- cific tolerance in juveniles of Coelotes terres- tris (Wider 1834) could be lengthened when there is an abundant supply of food. Similarly, Rypstra (1986) found that Achaearanea tepi- dariorum (C.L. Koch 1841), a solitary spe- cies, remained sociable longer when prey was abundant. Small and large spiders in the dyads did not differ in the number of hourly scans observed feeding. However it was noted that the larger spider usually fed before its smaller counter- part if a large size discrepancy was present. Perhaps by feeding first the larger spider gained more nutritionally by feeding until sa- tiation. This behavior could be seen as dom- ination, but the smaller spider did feed later and for the same span of time. No difference was seen in the number of hourly intervals observed feeding within the individuals in tet- rad groups. The most common feeding display in this treatment group was one spider feeding at a time. As expected, weight gain correlated posi- tively with hourly intervals observed feeding. Singletons, dyads, and tetrads were all ob- served feeding during the same number of hourly intervals. If there were some foraging advantage associated with feeding in groups, then individuals in groups of four should gain as much weight, or more, as individuals reared alone and spend less time feeding. Since this was not observed, there may be reasons (other than improved feeding efficiency) for social behavior to exist among young H. gigas. These benefits may include a reduced risk of predation, and the advantage of cooperatively seizing live prey. Tetrads of H. gigas spiderlings in this study exhibited cluster feeding only 6.9% of the time. Cluster feeding is a communal feed- ing behavior that involves the spiders hud- dling with their legs intertwined. This is an unusual behavior that has been documented in Aebutina binotata Simon 1892, a com- munal cribellate spider (Aviles 1993). Adult A. binotata females captured and communal- ly fed on large prey items such as cockroach- es and beetles; juveniles fed when the adults left the prey (Aviles 1993). Jantschke and Nentwig (2001) observed spiderlings of the mygalomorph spider, Ischnothele caudata feeding together on a prey item provided by the mother, but there was no mention of the specific cluster feeding behavior such as we observed in H. gigas. Ischnothele caudata ju- veniles will also cooperate in catching larger prey for up to 18 weeks. Reichling and Gutz- ke (unpubl.) found H. crassipes spiderlings clustering around food items caught by the mother. In our study there was only one instance of cannibalism in 18 groups: a spiderling in a tetrad killed its three siblings. Given the time span of the experiment and the number of group-reared H. gigas involved, we can tentatively conclude that cannibalism is rare in sibling groups of this species. The occur- rences of agonistic behaviors in general may have been undercounted due to the use of scan sampling, because probabilities of re- cording temporally short displays of hostility or facilitation are low. Although agonistic be- havior was observed, more lengthy focal ob- servations might have better documented these interactions. Agonistic displays were similar in incidence in tetrads and dyads but did not occur in higher frequency as expected in tetrads, where more spiders were forced to interact. Aggression was not more readily ob- served between similarly-sized spiders, con- trary to the group-living pholcid spider Hol- ocnemus pluchei (Scopoli 1763) where fights over prey were most intense between spiders of comparable size (Jakob 1994). Instead, ag- gression was positively correlated with the coefficient of variation of final weights, showing that replicates containing spiders with large size discrepancies either engaged in more aggressive displays or aggression led to large size discrepancies. This concurs with findings on the social spider Anelosimus ex- imius in which larger females commandeer prey captured by smaller females (Ebert 1998). Hysterocrates gigas spiderlings exhibited an unusual level of mutual tolerance, but this sociality did not apparently result in facilita- tion of feeding behavior, despite the distinc- tive cluster feeding posture we observed. Ta- rantulas, like all spiders, are generally cannibalistic beyond a short period of mutual toleration when young. Hysterocrates may be among the most sociable of theraphosid spi- ders. We have observed that H. gigas spider- lings in captivity will cohabit until several months of age. However, in a pilot study we conducted, H. gigas spiderlings that had been 330 THE JOURNAL OF ARACHNOLOGY first separated and then placed in social groups engaged in high levels of cannibalism, indi- cating that the suppression of cannibalism may depend on keeping the spiderlings in so- cial groups after hatching. Cohabitation of sib- ling groups has been observed in at least three other mygalomorph spider taxa: Nemesia ce~ mentaria (Buchli 1969), Heterothele darcheni (Darchen 1967) and Pamphobeteus sp. Po- cock 1901 (Cocroft & Rambler 1989). It re- mains to be seen how widespread this behav- ior is. For social behavior to evolve organisms must have something to share, in the case of spiders this is a web or retreat, as well as an abundance of prey (Shear 1970; Rypstra 1993; Leborgne et al. 1998; Jantschke & Nentwig 2001). Contrary to Jantschke and Nentwig’s (2001) claim that all social spiders must have a shared web for information transfer, very few mygalomorphs build webs. They construct burrows and in some cases the young will stay in the maternal burrow for extended periods of time (Buchli 1969). The burrow may promote sociality in the same way as a prey capture web. The in- creased level of sociality observed in this ta- rantula may result from the selective advan- tages accrued from sharing the deep maternal burrow and receiving protection from pred- ators and harsh environmental conditions. Hysterocrates exhibits a high level of mutual tolerance and even unique feeding behaviors associated with prey sharing, making it an unexpectedly social tarantula. However, we have shown that group size does not influ- ence the rate of growth. ACKNOWLEDGMENTS We thank M. Harless and R. Blatchford for assistance in collecting data. We thank S. Reichling, M. Hodge, R. West, M. Ashton and two anonymous readers for their comments on an earlier draft of this manuscript. And finally, we thank Dr. Bruce and Mrs. Janet Johnson for their donation to the J. H. Barrow Eield Station Research Eund, which made this re- search possible. LITERATURE CITED Aviles, L. 1993. Newly discovered sociality in the neotropical spider Aebutina binotata Simon (Dictynidae?). Journal of Arachnology 21:184- 193. Aviles, L. 1997. Causes and consequences of co- operation and permanent-sociality in spiders. Pp. 476-498. In The Evolution of Social Behavior in Insects and Arachnids. (J. C. Choe & B. J. Crespi, eds.). Cambridge University Press, Cam- bridge, England. Buchli, H.R. 1969. Hunting behavior in the Cteni- zidae. American Zoologist 9:175-193. Cocroft, R.B. & K. Hamblen 1989. Observations on a commensal relationship of the microhylid frog Chiasmocleis ventrimaculata and the bur- rowing theraphosid spider Xenesthis immanis in southeastern Peru. Biotropica 21:2-8. Curtis, J.T & J.E. Carrel. 1999. Social behavior by captive juvenile Kuknlcania hibernalis (Araneae: Filistatidae). Bulletin of the British Arachnolog- ical Society 1 1(6):241-246. Darchen, R. 1967. Biologic d’une mygale gabona- ise nouvelle: Macrothele darcheni Benoit (Ara- neida, Dipluridae). Biologia Gabonica 4:253- 257. Ebert, D. 1998. Behavioral asymmetry in relation to body weight and hunger in the tropical social spider Anelosimus eximius (Araneae, Theridi- idae). Journal of Arachnology 26:70-80. Jantschke, B. & W. Nentwig. 2001. Sub-social be- haviour in the diplurid Ischnothele caudata (Ar- aneae, Dipluridae). Bulletin of the British Arach- nological Society 12:12-16. Jakob, E.M. 1994. Contests over prey by group- living pholcids (Holocnemus pluchei). Journal of Arachnology 22:39-45. Krafft, B., A. Horel, & J.-M. Julita. 1986. Influence of food supply on the duration of the gregarious phase of a maternal-social spider, Coelotes ter- restris (Araneae, Agelenidae). Journal of Arach- nology 14:219-226. Leborgne, R., T. Cantarella, & A. Pasquet. 1998. Colonial life versus solitary life in Cyrtophora citricola (Araneae, Araneidae). Insectes Sociaux 45:125-134. Marshall, S.D. 1996. Tarantulas and Other Arach- nids. Barron’s Educational Series Inc., Haup- pauge, NY. Reichling, S.B. Sl W.H.N. Gutzke. Unpublished Manuscript. Maternal food-sharing by the Afri- can baboon spider Hysterocrates crassipes af- fects Juvenile growth. Rypstra, A.L. 1986. High prey abundance and a re- duction in cannibalism: the first step to sociality in spiders (Arachnida). Journal of Arachnology 14:193-200. Rypstra, A.L. 1993. Prey size, social competition, and the development of reproductive division of labor in social spider groups. The American Nat- uralist 142(5):868-880. Shear, W.A. 1970. The evolution of social phenom- ena in spiders. Bulletin of the British Arachnol- ogical Society l(5):65-76. VARRECCHIA ET AL.— GROUP SIZE AND GROWTH IN H, GIGAS 331 Smith, A.M. 1990. Baboon Spiders: Tarantulas of Africa and The Middle East. Fitzgerald Publish- ing, London. Uetz, G.W. & C. Hieber. 1997. Colonial web-build- ing spiders: Balancing the costs and benefits of group-living. Pp. 458-475. In The Evolution of Social Behavior in Insects and Arachnids. (J. C. Choe & B. J. Crespi, eds.). Cambridge Univer- sity Press, Cambridge, England. Manuscript received 31 July 2002, revised 22 Au- gust 2003. 2004. The Journal of Arachnology 32:332-335 SHORT COMMUNICATION A NEW SPECIES AND A NEW SYNONYMY IN THE SPINY ORB-WEAVER SPIDER GENUS MICRATHENA (ARANEAE, ARANEIDAE) Marcelo O. Gonzaga: Pos-Graduagao em Ecologia, Departamento de Zoologia, UNICAMP, Caixa Postal 6109, 13083-970, Campinas, SP, Brasil. E-mail: mogonz@unicamp.br Adalbert© J. Santos: Pos-Graduagao em Zoologia, Universidade de Sao Paulo. Laboratorio de Artropodes, Institute Butantan, Av. Vital Brasil 1500, 05503-900, Sao Paulo, SP, Brazil. E-mail: oxyopes@yahoo.com ABSTRACT. A new species, Micrathena ciciita, is described and illustrated based on females from southeastern Brazil. Plectana degeeri Walckenaer is synonymized with M. plana (C.L. Koch), based upon the original description. Keywords: Micrathena, Brazil, South America, systematics, new species The genus Micrathena Sundevall 1833 is a com- mon and conspicuous group of diurnal spiders en- demic to the Americas. They can be easily recog- nized by their spiny abdomen, vertical orb-webs with an open hub and their characteristic upside down position on the webs, with the abdomen in- clined horizontally. They differ from Gasteracantha Sundevall 1833 by the shape of carapace and from Chaetacis Simon 1895 by lacking spines or tuber- cles on the carapace behind the lateral eyes (Levi 1985). The genus was revised by Levi (1985) and comprises 104 species with a mostly Neotropical distribution. Of these, 33 are known only from fe- males and except for two notes describing males of previously known species (Bonaldo 1990; Lise 1995) and the synonymy of Thaumastobella niourei Mello-Leitao 1945 with Micrathena saccata (C.L. Kock 1836) (Scharff 1991), the systematics of the genus has remained unaltered since Levi’s revision. In this paper we describe a new species, Micrath- ena cicuta, of the kirbyi group (as defined by Levi 1985). This is the largest species group in this ge- nus and includes 45 species distributed from Cen- tral America to southern South America. Addition- ally, we synonymize M. degeeri (Walckenaer 1842) with M. plana (C.L. Koch 1836), another member of the kirbyi group, based upon the original descrip- tion. The specimens examined were deposited in the spider collection of the Instituto Butantan, Sao Pau- lo (I BSP) and Museu de Zoologia, Universidade de Sao Paulo (MZSP). The description format follows Levi (1985) and all measurements are in millime- ters. RESULTS Micrathena cicuta new species Figs. 1-5 Material examined. — Holotype female, Volta Redonda, Area de Relevante Interesse Ecologico Floresta da Cicuta, 22°3LS 44°07'W, State of Rio de Janeiro, Brasil, 18 February 2002, M.O. Gon- zaga (IBSP 36322). Paratype: 1 female, Resende, District of Serrinha do Alambari, 22°20'S 44°30'W, State of Rio de Janeiro, Brasil, E.F Ramos, 31 March 1996 (IBSP 27195). Other material exam- ined: BRASIL: Sao Paulo: Salesopolis, Esta^ao Biologica de Boraceia, May 2001, equipe BIOTA, 1 9 (IBSP 39799); Cotia, Reserva Florestal de Morro Grande, 26 March 2003, A. A. Nogueira et ak, 1 9 (IBSP 39794), 4 9 (IBSP, 39795-39798); 15 9 (MZSP). Etymology. — The specific name is a noun in ap- position taken from the type locality. Diagnosis. — Micrathena cicuta shares with Mi- crathena clypeata (Walckenaer 1805) a flat abdo- men with rounded sides, covered by several scler- otized disks and minute granules and with one anterior pair of spines. It is distinguished by the abdomen with 8 marginal pairs of black thorns (5 in M. clypeata) and by the absence of dimples on the carapace (Fig. 1). The epigynum differs by the narrower lateral lobes and median plate in ventral 332 GONZAGA & SANTOS— NEW SPECIES OF MICRATHENA 333 Figures 1-4. — Micrathena cicuta new species. 1. Female habitus, dorsal view. 2. Female epigynum, posterior view. 3. Ventral. 4. Lateral. Scale bars, 1 = 1.00 mm; 2-4 = 0.50 mm. and posterior views (Figs. 2, 3), the anteriorly notched transverse bar with a longer and posteriorly directed lobe (Fig. 3) and by the rounded bulge in lateral view (Fig. 4). Description. — Male.- Unknown. Female (holotype): Carapace orange, with a high thoracic region. Clypeus, chelicerae, labium, en- dites and sternum orange. Palpus and legs orange- brown, darker ventrally. Abdomen orange, lighter than carapace, with one anterior pair of spines over- hanging carapace and 8 pairs of marginal black thorns. Dorsum of abdomen flat with a slight pos- terior median longitudinal depression. Dorsal scler- otized disks and minute granules as in M. clypeata. Total length 8.6, carapace 2.8 long, 2.5 wide. First femur 2.3; patella and tibia 2.5; metatarsus 1.0; tar- sus 0.6. Second patella and tibia 2.3; third patella and tibia 1.4. Fourth femur 3.2; patella and tibia 2.3; metatarsus 1.6; tarsus 0.7. Distribution. — Known only from southeastern Brazil. This species seems to be closely related to M. clypeata, which occurs in Panama, northern South America and the Amazon Basin (Fig. 5). These two species display a disjunct distribution, occurring in tropical rainforest areas separated by central and northeastern Brazilian savanna vegeta- tions (cerrado and caatinga, Hueck 1972). Micrathena plana (C.L. Koch) Acrosoma planum C.L. Koch 1836:81, fig. 228. Micrathena plana: Levi 1985:509, figs. 311-325, map 6. Plectana degeeri Walckenaer 1842:174 (female ho- lotype from Suriname, lost) NEW SYNONYMY. Acrosoma degeeri: Butler 1873:425. Micrathena degeeri: Petrunkevitch 1911: 368; Plat- nick 2003. Remarks. — Micrathena degeeri was overlooked 334 THE JOURNAL OF ARACHNOLOGY Figure 5. — Geographic distribution of Micrathena clypeata (gray area, based on records from Levi 1985) and locality records for Micrathena cicuta new species (circles). by Levi (1985), and the type specimens are lost like so many of the species described by Walckenaer (1842). According to the original description (Wal- ckenaer 1842:174), M. degeeri possesses an oval- triangular abdomen with 12 spines: an anterior pair of small ones (described as “mediocre” by the au- thor), a pair of large and diverging posterior spines with two small ones on the base, one dorsal and one ventral. The sides of the abdomen were de- scribed as bearing two small spines. This descrip- tion matches Levi’s (1985) illustrations of Micrath- ena plana, a species distributed from Panama and the West Indies to Argentina, and the only species with 12 abdominal spines recorded from Suriname. As such, we here consider M. degeeri a junior syn- onym of M. plana. We are especially grateful to Angela M.F. Pache- co for the illustrations, and Cristina A. Rheims, Ri- cardo Pinto da Rocha, and Andre A. Nogueira for providing additional specimens for this study. We are also indebted to Fundagao CSN and IB AM A for allowing our studies in the ARIE Floresta da Cicuta. Antonio D. Brescovit, C.A. Rheims and two anonymous reviewers are acknowledged for helpful suggestions on the manuscript. This study was fi- nanced by FAPESP doctoral fellowship grants (Proc. 99/06089-4 to M.O. Gonzaga and 99/05659-8 to A.J. Santos). GONZAGA & SANTOS— NEW SPECIES OF MICRATHENA 335 LITERATURE CITED Bonaldo, A.B, 1990. Descrigao do macho de Mi~ crathena kirbyi (Perty, 1833) (Araneae, Aranei- dae). Iheringia (ser. ZooL) 70:89-91. Butler, A.G. 1873. A list of the spiders of the genus Acrosoma. Proceedings of Zoological Society of London 1873:420-429. Koch, C.L. 1836. Die Arachniden. Nurnberg. Hueck, K. 1972. As Florestas da America do Sul: ecologia, composigao e importancia ecoeomica. Poligono/Editora da Universidade de Brasilia, Sao Paulo. Levi, H.W. 1985. The spiny orb-weaver genera ME crathena and Chaetacis (Araneae: Araeeidae). Bulletin of the Museum of Comparative Zoology 150:429-618. Lise, A. A. 1995. Description of the male of ME crathena peregrinatorum (Holmberg, 1883) (Ar- aneae, Araneidae). Revista Brasileira de Ento- mologia 39:221-223. Mello-Leitao, C.E 1945. Tres novas especies de Gasteracanthinae e notas sobre a subfamflia. An- ais da Academia brasileira de Ciencias 17:261- 267. Petruekevitch, A. 1911. A synonymic index-cata- logue of spiders of North, Central and South America with all adjacent islands, Greenland, Bermuda, West Indies, Terra del Fuego, Gala- pagos, etc. Bulletin of American Museum of Natural History 29:1-791. Platnick, N.L 2003. The world spider catalog, ver- sion 3.5. American Museum of Natural History, on line at Scharff, N. 1991. On the synonymy of Thaumas- tobella mourei Mello-Leitao and lldibaha albom- aculata Key selling (Araneae, Araneidae). Jour- nal of Arachnology 19:155-156. Walckenaer, C.A. 1842. Histoire naturelle des In- sectes, Apteres. Vol. 2. Librairie Encyclopedique de Roret, Paris. Manuscript received 27 May 2003, revised 2 Sep- tember 2003. 2004. The Journal of Arachnology 32:336-340 SHORT COMMUNICATION COLOR DIMORPHISM IN ADULTS AND JUVENILES OF BUITINGA SAFURA (ARANEAE, PHOLCIDAE) Bernhard A. Huber: Zoological Research Institute and Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany. E-mail: b.huber.zfmk@uni-bonn.de Sascha Hopf: Konigsberger Str. 18, 53840 Troisdorf, Germany ABSTRACT. We document the hrst case of a color dimorphism in a pholcid spider. Males, females and juveniles of Buitinga safura Huber 2003 either have large black spots on the abdomen or no spots, with no intermediates. At the same time, this species shows sexual dimorphism (brown prosomal marks present in males only) and continuous prosomal pattern variation in males, females and juveniles. The abdominal pigment is located in the hypodermis. Keywords: Color dimorphism, polymorphism, pattern variation, Pholcidae Intrasexual polymorphism (discontinuous indi- vidual variation among members of the same sex, in the same life stage, within a population) is con- sidered a key phenomenon for the study of basic evolutionary concepts (e.g., West-Eberhard 1989; Gould 1989; Eberhard & Gutierrez 1991; Mayr & Ashlock 1991; Emlen 1994). Color polymorphisms, being easily visible, are among the best studied and considerable progress has been made introducing spiders as possible model arthropods with which to study the evolutionary processes working on visi- ble, intraspecific variation. Spectacular examples include the candy-striped spider, Enoplognatha ova- ta (Clerck), and the Hawaiian happy-face spider, Theridion grallator Simon, but many cases have been described in various spider families. Extensive reviews on spider coloration and polymorphism have been published recently by Oxford & Gillespie (1998) and Oxford (1999). We present here the first case of a color poly- morphism in the spider family Pholcidae. A large series of Buitinga safura Huber 2003 was collected in the Uzungwa Mountains, Iringa Province, Tan- zania (for details of study site, see Sprensen et al. 2002), by an expedition of the Smithsonian Insti- tution in Washington, D.C. and the Zoological Mu- seum of Copenhagen in May 1997. The material studied herein is deposited in the National Museum of Natural History (Washington, D.C., U.S.A.). Specimens were collected from litter and logs, low vegetation, understory and canopy, and transferred to 70% ethanol (Sprensen et al. 2002). The present analysis is restricted to the largest subsample, i.e. to the 1 139 specimens collected from the understo- ry, but inspection of the other material suggests that the patterns described below do not differ signifi- cantly among habitats. We measured nine traits (Figs. 1-4) with a measuring grid in the ocular of a Nikon SMZ-U dissecting microscope and as- sessed prosomal pattern variation in a qualitative way (Figs. 5-8). Tables 1 and 2 give the sample sizes, means, ranges, standard deviations, coeffi- cients of variation, significance values of Kolmo- gorov-Smirnov tests for normal distribution and es- timates of measurement error for all measured traits. Figures 1-4. — Illustrations of characters mea- sured. 1. Prosoma and abdomen, dorsal view; a = abdomen length, e = eye distance, pm = posterior mark on carapace, si, s2 = abdominal spot 1 and 2 lengths. 2. Tibia, lateral view; tib 1,3 = tibia 1 and tibia 3 lengths. 3. Right male chelicera, lateral view; chel: chelicera length. 4. Left procursus, pro- lateral view; proc = procursus length. 336 HUBER & HOPE— COLOR DIMORPHISM IN A PHOLCID SPIDER 337 Figures 5--10. — Photographs of six adult Buitinga safura specimens showing some of the color variation described: abdominal spots present (5-8) vs. absent (9-10), posterior mark on carapace present (6-7) vs. absent (5, 8-10), brown bands on carapace present (6, 8, 10) vs. absent (5, 7, 9), and the four arbitrarily defined degrees of lateral prosomal patterns: a (only black lines), b (black lines plus brown bands), c (black lines plus three pairs of black spots), d (black lines plus brown bands plus three pairs of black spots). From 341 adult males in the sample, 20 had two pairs of spots on the abdomen, two had only one (the posterior) pair of spots. These 22 spotted males were all measured. From the remaining 319 spotless males, 23 were randomly selected and included in the quantitative analysis, resulting in a total of 45 males measured. Histograms of male spot lengths clearly indicate that these are not cases of contin-= uous variation (Figs. 11, 12). All other traits mea- sured (with the exception of the posterior mark on the carapace, see below) show unimodal distribu- tions that are not significantly different from normal distributions (Table 1). All specimens are consid- ered conspecific because those characters that in pholcids differ most among species (procursus, bulb, cheliceral armature; see Huber 2003) were virtually identical. From a scatter between two char- acters with high interspecific variation (procursus, chelicerae), it is evident that spotted (o) and spot- less (+) males occur at any sizes of these characters (Fig. 13). Also, there was no correlation between overall size and abdominal pattern (Fig. 14; t-test calculated for tibia 1 length, tibia 3 length, and eye distance: all P > 0.05). Ordinary least squares (OLS) regressions of log-transformed characters were calculated for all traits on eye distance as an indicator of body size (for justification of method see Eberhard et al. 1999). As in the comparative study by Eberhard et al. (1998), legs and other non- genitalia had relatively high slopes (tibia 1 length Table 1. — ^Male characters measured, with sample sizes («), means, ranges, standard deviations (SD), coefficients of variation (CV), significance values of Kolmogorov-Smirnov tests for normal distribution (KS), and estimates on measurement error. Character n Mean (mm) Range (mm) SD CV Measurement KS error (± m,m) Tibia 1 length 39 5.65 5.00-6.13 0.286 5.1 0.81 0.07 Tibia 3 length 44 2.48 2.20-2.73 0.109 4.4 0.47 0.03 Abdomen length 45 1.90 1.58-2.25 0.166 8.7 0.24 0.03 Eye distance 44 0.56 0.52-0.60 0.018 3.2 0.21 0.01 Chelicera length 44 0.60 0.53-0.63 0.024 4.2 0.62 0.01 Procursus length 43 0.53 0.51-0.56 0.012 2.4 0.48 0.01 Abdominal spot 1 20 0.48 0.38-0.60 0.061 12.7 0.00 0.03 Abdominal spot 2 22 0.24 0.15-0.35 0.042 17.4 0.00 0.03 Carapace posterior mark 7 0.26 0.05-0.38 0.117 44.3 0.00 0.03 338 THE JOURNAL OF ARACHNOLOGY Table 2. — Female characters measured, with sample sizes {n), means, ranges, standard deviations (SD), coefficients of variation (CV), significance values of Kolmogorov-Smirnov tests for normal distribution (KS), and estimates on measurement error. Character n Mean (mm) Range (mm) SD CV KS Measurement error (± mm) Tibia 1 length 50 4.75 4.37-5.23 0.196 4.1 0.70 0.07 Tibia 3 length 54 1.98 1.87-2.17 0.070 3.5 0.19 0.03 Abdomen length 56 1.75 1.38-2.18 0.157 9.0 0.87 0.03 Abdominal spot 1 29 0.49 0.35-0.73 0.084 17.3 0.00 0.03 Abdominal spot 2 31 0.21 0.13-0.28 0.037 17.7 0.00 0.03 Carapace posterior mark 4 0.20 0.13-0.28 0.061 30.6 0.00 0.03 = 1.00; tibia 3 length = 0.79; chelicerae length = 0.73; all P < 0.001), while procursus length had a much lower slope (0.38; P < 0.001), as is usual for genitalia. This trend remained when spotted and un- spotted individuals were analyzed separately, but there was considerable variation among slopes of spotted vs. unspotted males (probably due to small sizes of subsamples). The slope of abdomen length on eye distance was non-significant. Male prosomal pattern variation was also sub- stantial. However, this variation was continuous and not dimorphic. We arbitrarily defined four types within the continuum of lateral prosomal patterns (Figs. 5-8), but found no significant correlation of these with body size (Fig. 15) or abdominal pattern (Fig. 16). In seven (out of 341) males, there was a large black mark posteriorly on the carapace (Fig. 6). This trait may be dimorphic too, but sample size is obviously too small. All of these males also had abdominal spots, but two of them had only the pos- terior pair. In females we measured the same traits except for eye distance, procursus length and chelicera length. From the 385 adult females in the sample, 29 had both pairs of abdominal spots, two had only one (the posterior) pair, all others were spotless. 30 7 Figures 11,1 2. — Histograms showing the bimodal distributions of abdominal spot sizes in males. Note that the left bars indicating spotless specimens represent only a small fraction of the more than 300 spotless males in the original sample. HUBER & HOPE— COLOR DIMORPHISM IN A PHOLCID SPIDER 339 + = spotless) in males with different sizes of chelicerae and procursi (13) and in males and females of different overall size as indicated by leg length (14). Fig. 13 strongly supports conspecificity of spotted and spotless males, while Fig. 14 shows that there is no correlation between abdominal spottedness and size. Lateral pattern on carapace Figures 15, 16. — 15. Scatter diagram showing the distribution of lateral prosomal patterns in males and females of different sizes: pattern “a” (Fig. 5; represented by squares) occurs in males and females but is rare in males; patterns “b” and “d” (Figs. 6 and 8; represented by circles) occur only in males; pattern “c” (Fig. 7; represented by crosses) occurs in both sexes. 16. Bar diagram showing that abdominal spots occur in males with all different kinds of prosomal patterns. Sample size is too small to judge the signif- icance of the increase in cases of abdominal spots from a-d. 340 THE JOURNAL OF ARACHNOLOGY Figure 17. — Semithin section of a female abdo- men in the area of a spot, showing the location of the pigment in the hypodermis; c = cuticle, m = subcuticular muscle layer, p = pigment. Scale line: 25 (xm. From these, 25 randomly chosen individuals were measured resulting in a total of 56 measured fe- males. Abdominal spot sizes in females were sim- ilar to those in males (see Tables 1 and 2) and the proportions of spotted vs. unspotted specimens was not significantly different in males vs. females (chi square = 0.684, P — 0.408, 1 df). However, females never had brown marks on the prosoma, i.e. they showed only two of the four lateral prosomal pat- terns shown in Figs. 5-8. Therefore, in addition to the abdominal intrasexual dimorphism there is also an (inter)sexual dimorphism. Posterior black marks occurred in four females and all of these also had abdominal spots, but two of them only the posterior pair. The epigyna of all females were indistinguish- able. In juveniles we only counted the numbers of spotted and spotless specimens (19 vs. 394). The percentage of spotted individuals was similar to that in adults (chi square == 4.022, P = 0.134, 1 df). Most juveniles were late or penultimate instars. Pro- somal pattern variation in juveniles appeared simi- lar to that in females, but most juveniles had only the black lines (cf. Fig. 5) and lateral spots, if pre- sent, were usually very weak. Posterior black marks on the carapace were not seen in juveniles. The abdominal pigment is located in the hypo- dermis: removal of the digestive tract left the spots intact, but after treatment with NaOH they could be removed easily from the cuticle using a brush. This result was confirmed by preparation of semithin sections (Fig. 17). By comparison with other spider pigments (Oxford & Gillespie 1998), the location suggests it is an ommochrome. There is no evidence that preservation in ethanol has had any effect on the abdominal spots: they are either deep black or entirely absent. Little can be said beyond these ba- sic facts. The truly interesting questions remain to be answered: is the polymorphism genetically de- termined? Is it selectively maintained, and if yes, by which selective forces? What are the costs of producing spots, if any? How many alleles contrib- ute to the polymorphism, and which morph is dom- inant? We are grateful to Jonathan Coddington for the loan of the material treated in this paper and to Geoff Oxford, Gail Stratton and two anonymous referees for valuable comments on a previous ver- sion of the manuscript. LITERATURE CITED Eberhard, W.G. & E.E. Gutierrez. 1991. Male di- moiphisms in beetles and earwigs and the ques- tion of developmental constraints. Evolution 45: 18-28. Eberhard, W.G., B.A. Huber, S. Rodriguez, R.L., Briceno, R.D. Salas, 1. & V. Rodriguez. 1998. One size fits all? Relationships between the size and degree of variation in genitalia and other body parts in twenty species of insects and spi- ders. Evolution 52(2):415-43 1. Eberhard, W.G., B.A. Huber, & R.L. Rodriguez. 1999. Don’t forget the biology: a reply to Green. Evolution 53(5): 1624-1627. Emlen, D.J. 1994. Environmental control of horn length dimorphism in the beetle Onthophagus aciiminatiis (Coleoptera: Scarabaeidae). Proceed- ings of the Royal Society, London B 256:131- 136. Gould, S.J. 1989. A developmental constraint in Cerion, with comments on the definition and in- terpretation of constraint in evolution. Evolution 43:516-539. Huber, B.A. 2003. High species diversity in one of the dominant groups of spiders in East African montane forests (Araneae: Pholcidae: Buitinga n. gen., Spennophora Hentz). Zoological Journal of the Linnean Society 137:555-619. Mayr, E. & P.D. Ashlock. 1991. Principles of Sys- tematic Zoology. McGraw-Hill, Inc. Oxford, G.S. & R.G. Gillespie. 1998. Evolution and ecology of spider coloration. Annual Review of Entomology 43:619-643. Oxford, G.S. 1999. Evolutionary significance of colour variation in arthropods: lessons from the spiders. Boletin de la Sociedad Entomologica Aragonesa 26:619-635. Sprensen, L.L, J.A. Coddington & N. Scharff. 2002. Inventorying and estimating subcanopy spider diversity using semiquantitative sampling methods in an afromontane forest. Environmen- tal Entomology 3 1(2):3 19-330. West-Eberhard, M.J. 1989. Phenotypic plasticity and the origin of diversity. Annual Review of Ecology and Systematics 20:249-278. Manuscript received 28 April 2003, revised 17 Sep- tember 2003. 2004. The Journal of Arachnology 32:341-344 SHORT COMMUNICATION PARDOSA MILVINA (ARANEAE, LYCOSIDAE) SPIDERLING MOVEMENT IN THE PRESENCE OF CONSPECIFIC AND HETEROSPECIFIC SILK AND EXCRETA Matthew H. Persons and Erin C. Lynam: Biology Department, Susquehanna University, 514 University Avenue, Selinsgrove, 17870, PA, USA ABSTRACT. Adult females of the wolf spider Pardosa milvina (Hentz 1844) are known to show adaptive antipredator responses in the presence of chemical cues (silk and excreta) from the larger co- occurring wolf spider Hogna helluo (Walckenaer 1837). We tested if the presence of H. helluo cues affected P. milvina spiderling dismounts from their mothers. Immediately after females opened their egg sacs, we counted offspring and placed spiderling-caiTying females on one of three experimental substrates: 1) container previously occupied for 24 h by an adult conspecific female, 2) container previously occupied for 24 h by a juvenile H. helluo equal in mass to an adult F. milvina, or 3) a control container devoid of either cue. We then measured the proportion of spiderlings that dismounted from their mothers over a six- day period. Spiderling dismounts peaked by day three, after which spiderlings tended to return to their mother. During day one, significantly fewer spiderlings were dismounted from the mother in containers previously occupied by a juvenile H. helluo compared to other treatments. There was no significant difference in dismounts among treatments during days 2-6. Since spiderlings were maximally dismounted by day three, we suggest that spiderlings may tend to disperse into areas with fewer H. helluo. Keywords: Spiderlings, lycosid, dispersal, chemical cue, Hogna helluo The wolf spider Pardosa milvina (Hentz 1844) and its larger intraguild predator, Hogna helluo (Walckenaer 1837) are common throughout agri- cultural systems in the Midwestern United States (Marshall & Rypstra 1999). Recent studies show that predator-prey interactions among these species are mediated in part through each other’s silk drag- lines and excreta (Persons & Rypstra 2000; Persons et al. 2001). Adult female P. milvina respond to chemical cues (silk and excreta) from adult and ju- venile H. helluo with a suite of antipredator behav- iors including substratum avoidance, reduced activ- ity and vertical movement (Persons et al. 2001, 2002; Persons & Rypstra 2001). In contrast, adult female P. milvina are unresponsive to conspecific female silk and excreta (Persons et al. 2001, 2002). Female wolf spiders typically carry their egg sac by attaching it to their spinnerets for a period of several days to more than two months and after opening the egg sac, continue to transport spider- lings on their dorsum for a period of several days to a week or more (Fuji! 1976). Spiderlings then disperse primarily via asynchronous dismounting from their mother as she moves through the envi- ronment (Fujii 1976) and secondarily via aerial dis- persal (Click 1939; Richter 1970). Juveniles may also periodically dismount and remount their moth- ers prior to permanent dispersal to drink or repo- sition themselves for more efficient transport (Ea- son 1964; Higashi & Rovner 1975). Pardosa milvina spiderlings may benefit substan- tially by dismounting their mother when in micro- habitats that are devoid of H. helluo and their as- sociated chemical cues. Although predator-induced reductions in activity increase survival of P. milvina when in the presence of a live H. helluo (Persons et al. 2001, 2002; Barnes et al. 2002), these defen- sive behaviors are costly and contribute to reduced foraging and reproductive success (Persons et al. 2002). Also, under some circumstances, H. helluo are attracted to substrates previously occupied by adult female P. milvina (Persons & Rypstra 2000) which may further increase predation risk to P. mil- vina spiderlings. Here we tested if the presence of silk and excreta from juvenile H. helluo signifi- cantly affected the timing of juvenile P. milvina dis- mounts from their mothers. Sixty-eight mated adult female P. milvina were field caught in late May and early June, 2000 within soybean fields on Susquehanna University property adjacent to campus (Selinsgrove, Snyder County, Pennsylvania). All field-collected females were ei- ther carrying egg sacs or produced them shortly af- ter being caught. Spiders were individually main- 341 342 THE JOURNAL OF ARACHNOLOGY tained with their egg sacs in white plastic containers with transparent lids (9 cm d, 7 cm h). A small vial lid was placed on the bottom of the container with a few drops of water to serve as a source of mois- ture and humidity. All spiders were kept at room temperature (23 — 25 °C) with a 13L:11D photope- riod. Females were given constant access to fruit flies {Drosophila melanogaster (Meigen 1 830) as a food source and checked daily for spiderling emer- gence from the egg sac. Because filial or sibling cannibalism is possible among lycosids, we chose to count spiderlings immediately after egg sac emergence to insure an accurate measure of clutch size. After emergence, spiderlings were gently re- moved from the mother with a soft-bristled paint- brush, counted, and allowed to climb back on their mother. The females with offspring were then ran- domly assigned to one of three substrate treatments: 1) a container that previously held a juvenile H. helluo for 24 h that was equal in mass to an adult female P. milvina {n — 25); 2) a container that pre- viously held an adult conspecific female P. milvina (without an egg sac), {n = 22) for 24 h; or, 3) a blank control container devoid of either cue {n = 21). Pardosa milvina and H. helluo used for gen- erating chemical cues were maintained on an ad li- bitum diet of fruit flies {D. melanogaster) prior to being placed in their respective treatment contain- ers. Hogna helluo and P. milvina were not fed dur- ing the time they were used to deposit silk and ex- creta nor were test P. milvina allowed to feed during the trial period. The deposited silk and ex- creta used as test substrates was not renewed for the duration of the experiment. Female test spiders were initially checked 24 h after being placed on their respective substrate treatments and again ev- ery 24 h afterwards for six consecutive days. All P. milvina with spiderlings were provided with a small inverted vial cap filled with water during the entire test period. The total number and proportion of spiderlings that had dismounted from their moth- er was recorded for each replicate. Only a spider- ling that had no direct physical contact with its mother was considered dismounted. Egg sac clutch sizes of P. milvina for all treat- ments varied between 1-101 spiderlings (mean = 31.8 ± S.E. 2.18 offspring; median = 30.5 off- spring). Therefore we used the proportion of dis- mounted spiderlings/egg sac as the dependent var- iable in our analysis. We angular transformed our proportions to conform to assumptions of normality and performed a repeated-measures two-way AN- OVA with chemical cue treatment (fixed effect) and day (random effect) as factors (as in Sih & Mc- Carthy 2002). Peak spiderling dismounts occurred three days post-emergence (mean 78.3% for all treatments combined)(Fig. 1). By day six of post-emergence, the mean proportion of dismounted spiderlings re- turned to similar levels observed during the first day. Substrate type had a significant effect on spi- derling dismounts (F265 = 3.69; P = 0.0301). There was also a significant day effect (Fg 55 = 22.22; P < 0.0001). However, there was no significant day by treatment interaction (F52 = 1.22; P = 0.276). To better understand dismounting patterns across chemical cue treatments but within days, we per- formed six a posteriori multiple comparison tests (Tukey test for unequal sample sizes) for ANOVAs as described by Zar (1984). Based on the results of the Tukey tests, signifi- cant substrate effects were attributable solely to dif- ferences in spiderling dismounts during the first day of emergence. Females placed in containers with H. helluo chemical cues had an average of only 28% (mean 8.76 ± S.E. 2.36 spiderlings) of their clutch dismounted by day one compared to 41% (mean 13.23 ± S.E. 3.20 spiderlings) and 55% (13.71 ± S.E. 2.44 spiderlings) for the P. milvina and control treatments respectively. During the first day of emergence, a significantly lower proportion of spi- derlings dismounted in the H. helluo cue treatment than either the P. milvina treatment or blank con- trol. However, the blank control and P. milvina treatments were not significantly different from each other. Proportion of spiderling dismounts among treatments for all other days was not statis- tically significantly different based on Tukey post- hoc comparisons. Other studies of spiderling dispersal times among Pardosa species have found ranges of 2-7 days for P. milvina (Montgomery 1903), 4-8 days for P. laura (Karsch, 1879)(Fujii 1976), and 3-7 days for P. t-insignita (Bosenberg & Strand 1906)(Fujii 1976). None of these studies noted remounting be- havior among spiderlings but dismounting and re- mounting for purposes of water seeking are known to occur among lycosids (Higashi & Rovner 1975). Since spiderlings were not marked, it was not pos- sible to determine the frequency of remounting by individual spiders. We noted that mounted spider- lings would frequently crawl down one of the moth- er’s legs and lightly tap at the ground. Afterwards they would either climb back onto the mother’s dor- sum, or dismount entirely. This behavior suggests a mechanism by which spiderlings may sample the substratum directly prior to dismounting. We also noted that dismounted spiderlings tended to re- mount their mother only after direct physical con- tact rather than showing directional orientation and approach toward the mother from other parts of the container. These observations indicate that re- mounting tends to be incidental with further mater- nal contact rather than a directed response from the spiderling at a distance. The differences in spiderling dismounts between treatments were modest, yet there was a signifi- cantly smaller proportion dismounted from their PERSONS & LYNAM— CHEMICAL CUES AND WOLF SPIDERLING DISPERSAL 343 0 1 2 3 4 5 6 7 Days since spiderling emergence Figure. — Mean proportion of spiderlings dispersed from their mothers (± S.D.) while on substrates previously occupied by a conspecific female P. milvina for 24 h (Pardosa cues), a juvenile H. helluo (Hogna cues), or a control substrate not previously occupied (Blank). mothers in the H. helluo treatment on day one of the experiment. In our study we did not renew the silk and excreta over the six-day experimental pe- riod. Previous studies indicate that P. milvina re- sponse to H. helluo cues becomes reduced over time due to the age of the stimulus (Barnes et al. 2002) rather than habituation to the cues (Persons et al. 2002). Adult females have a strong response toward fresh silk and excreta deposited by an adult H. helluo less than 24 hours earlier but are less responsive to one week-old H. helluo cues (Barnes et al. 2002). Results observed here are consistent with Barnes et al. (2002) in that the effect of H. helluo cues appeared to have diminished over the duration of the experiment and may have contrib- uted to the significant response during the first day but not on subsequent days. Given the high activity level of P. milvina (Walk- er et al. 1999), we propose that even small differ- ences in spiderling dismount frequencies across treatments may translate into favorable non-random site dispersal with respect to the presence of H. hel- luo. For now it remains unclear the extent to which variation in spiderling dismounts are attributable to spiderlings directly responding to predator cues or indirectly via subtle changes in the mother’s behav- ior in the presence of these cues. However, spider- ling dispersal via other mechanisms should be con- sidered as well. The prevalence of aerial ballooning or other forms of secondary dispersal in P. milvina 344 THE JOURNAL OF ARACHNOLOGY is unknown, but has been observed in other species in the genus (Richter 1970) and may further allow spiderling avoidance of areas with higher predation risk. ACKNOWLEDGMENTS We thank J.C. Owens for his help collecting and maintaining spiders used for this study. This research was funded in part through NSF grant C-RUI DBI- 0216776 (for M. Persons) and C-RUI DBI 0216947 (for A. Rypstra). Further support was received through a Susquehanna Research Partners stipend for E. Ly- nam. A male and female each of P. milvina and H. helluo have been deposited at the Denver Museum of Nature and Science as voucher specimens of this study. LITERATURE CITED Barnes, M.C., M.H. Persons & A.L. Rypstra. 2002. The effect of predator chemical cue age on an- tipredator behavior in the wolf spider Pardosa milvina (Araneae: Lycosidae). Journal of Insect Behavior 15:269-281. Eason, R.R. 1964. Maternal care as exhibited by wolf spiders (lycosids). Proceedings of the Ar- kansas Academy of Sciences 18:13-19. Fujii, Y. 1976. Pulli-carrying behaviour in wolf spi- ders (Lycosidae, Araneae). Bulletin of Nippon Dental University, General Education 5:144- 151. Higashi, G.A. & J.S. Rovner. 1975. Post-emergent behaviour of juvenile lycosid spiders. Bulletin of the British Arachnological Society 3:1 13-1 19. Glick, P.A, 1939. The distribution of insects, spi- ders, and mites in the air. United States Depart- ment of Agriculture Bulletin 673:1-150. Marshall, S.D. & A.L. Rypstra. 1999. Patterns in the distribution of two wolf spiders (Araneae: Lycosidae) in two soybean agroecosystems. En- vironmental Entomology 28:1052-1059. Montgomery, T.H. 1903. Studies on the habits of spiders particularly those of the mating period. Proceedings of the Academy of Natural Sciences Philadelphia 55:59-149. Persons, M.H. & A.L. Rypstra. 2000. Preference for chemical cues associated with recent prey in the wolf spider Hogna helluo (Araneae: Lycosidae). Ethology 106:27-35. Persons, M.H. & A.L. Rypstra. 2001. Wolf spiders show graded antipredator behavior in the pres- ence of chemical cues from different sized pred- ators. Journal of Chemical Ecology 27:2493- 2504. Persons, M.H., S.E. Walker, A.L. Rypstra, & S.D. Marshall. 2001. Wolf spider predator avoidance tactics and survival in the presence of diet-as- sociated predator cues (Araneae: Lycosidae). An- imal Behaviour 61:43-51. Persons, M.H., S.E. Walker, & A.L. Rypstra. 2002. Fitness costs and benefits of antipredator behav- ior mediated by chemotactile cues in the wolf spider Pardosa milvina (Araneae: Lycosidae). Behavioral Ecology 13:386-392. Richter, C.J.J. 1970. Aerial dispersal in relation to habitat in eight wolf spider species {Pardosa, Ar- aneae, Lycosidae). Oecologia 5:200-214. Sih, A., & TM. McCarthy. 2002. Prey responses to pulses of risk and safety: testing the risk alloca- tion hypothesis. Animal Behaviour 63:437-443. Walker, S.E., Marshall, S.D., Rypstra, A.L., & D.H. Taylor. 1999. The effects of hunger on locomo- tory behaviour in two species of wolf spider (Ar- aneae: Lycosidae). Animal Behaviour 58:515- 520. Zar, J.H. 1984. Biostatistical analysis. 2"^^ edition. Prentice-Hall, New Jersey. Manuscript received 17 March 2003, revised 27 August 2003. 2004, The Journal of Arachnology 32:345-348 SHORT COMMUNICATION DISTRIBUTION OF SPIDERS ON DIFFERENT TYPES OF INFLORESCENCES IN THE BRAZILIAN PANTANAL Andrea Lucia Teixeira de Souza and Erica de Souza Modena: Depto de Biologia/ CCBS — Setor Ecologia, Universidade Federal de Mato Grosso do Sul, CP 549, CEP 79070-900, Campo Grande, MS, Brazil. E-mail: altsouza@nin.ufms.br ABSTRACT. Reproductive stems add complexity to vegetation, thereby increasing the range and quality of microhabitats available for arthropods. In this study, we evaluated whether variation in inflorescence characteristics influenced spider distribution. We compared spider guild structure among inflorescences of three herbaceous plant species, Melanthera latifolia, Conyza bonariensis and Eupatorium hecatanthum (Asteraceae), and between inflorescences of C bonariensis in two different phenological stages, flower buds and opened flowers. Total spider abundance was higher on M. latifolia, intermediate on E. hecatan- thum, and lower on C bonariensis. Ambush spiders were more abundant on M. latifolia than on the other plant species, while the abundance of hunting spiders did not differ among plant species. Also, spiders recorded on M. latifolia were larger than those on both E. hecatanthum and C. bonariensis. However, ambush spiders were smallest on M latifolia, while hunting spiders on E. hecatanthum were larger than on the other plant species. The number of spiders on inflorescences with flower buds did not differ from those with opened flowers, but ambush spiders on inflorescences with opened flowers were larger than those on inflorescences with flower buds. Our results with different inflorescence types support the hy- pothesis that differences on microhabitat structure influence distribution of spiders. Keywords: Flower-dwelling spiders, Asteraceae, habitat structure Habitat structure can influence the abundance, di- versity and size distribution of spiders (Scheidler 1990; Evans 1997), since it is related to prey abun- dance, availability of refuges from predators and favorable microclimate conditions (Gunnarsson 1996; Halaj et al. 1998). Some studies have shown that the added complexity provided by higher den- sities of leaves and second order branchlets can re- sult in increased abundances and mean body sizes of some species of plant-dwelling spiders (Hatley & MacMahon 1980; Gunarsson 1990; Halaj et al. 2000). Inflorescences attract large numbers of herbivo- rous and pollinating insects due to the availability of pollen, nectar and edible tissues. The abundance of potential prey on plant reproductive stems can influence the assemblage of spiders that visit inflo- rescences (e.g., Morse & Fritz 1982; Neetwig 1993). In addition, the presence of inflorescences add another dimension to plant architecture by changing microclimate conditions and availability of refuges from predators. Structural characteristics of inflorescences such as branch size, texture, num- ber and size of leaves and flowers, and the arrange- ment of the biomass in space vary both among plant species and between inflorescences in distinct phe- nological stages (opened flowers vs. flower buds) within a plant species. However, few studies have considered the use of flowers by spiders, and pat- terns of spider distribution on inflorescences of dif- ferent plant species and/or in distinct phenological stages within the same plant species, are still ob- scure. In this study, we evaluated differences in the abundance and size distribution of crab spiders and hunting spiders among inflorescences of three plant species, and between reproductive stems in differ- ent phenological stages within the same plant spe- cies. This study was carried out in November 2000 at the Miranda-Abobral subregion of the Pantaeal do Mato Grosso, Central Brazil (19°34'S:57°00'W). The study area consists of natural forest fragments and gallery forests. These fragments have variable sizes, and are surrounded by seasonally flooded fields. Spiders were sampled from three species of herbaceous plants (Asteraceae), common at the edge of gallery forests on the riverside of Rio Mi- randa. The plant species sampled differed in several inflorescence characteristics: Melanthera latifolia (Gardn.) has a 40cm long inflorescence, with few (7.33 ± 1.53, mean ± SD; n = 10) yellow flowers, each one with a corolla diameter of 6.5 ± 0.87cm 345 346 THE JOURNAL OF ARACHNOLOGY Table 1. — Results of one-way ANOVAs and multiple comparisons tests comparing the number and body size of spiders (means ± standard errors) in different guilds (ambush or hunter) on inflorescences from three species of Asteraceae at the edge of a gallery forest. Similar letters connect means that did not differ (Tukey’s HSD, P > 0.05). Spiders M. latifolia C. bonariensis hecatanthum F P Number of spiders per inflorescence branch Total spiders 11.7 ± l.U 4.7 -1- 3.D 8.0 -i- 1.7"-b 8.073 0.020 Ambushers 9.3 ± 0.4" 2.0 -h 0.7” 4.3 -h 2.0^ 13.069 0.007 Hunters 1.3 ± 1.1 1.7 H- 1.5 3.7 -1- 0.8 1.792 0.245 Body size of spiders (mm) Total spiders 1.26 ± 0.07" 1.54 0.15" 3.31 -H 0.82^ 10.705 0.010 Ambushers 1.20 ± 0.07" 1.62 + 0.1 Qb 1.82 -f- 0.12b 12.771 0.007 Hunters 1.87 ± 0.13" 1.87 -+- 0.13" 4.41 -h 0.58b 21.081 0.008 (n = 10); Conyza bonariensis (L.) Cronq. has a 48cm long inflorescence, with many (33.0 ± 3.60; n =10) white flowers (corolla diameter = 1.30 ± 0.10cm); Eupatorium hecatanthum (DC.) Bak.has an inflorescence 1 2cm long, purple flowers (corolla diameter = 3.47 ± 0.15cm) and an intermediate number of flowers per inflorescence (22.67 ± 2.52) compared to the other two species. All plant species occurred together within the patches in the sam- pling sites. They grew intertwined, so that spiders could move from one species to another without spatial barriers. Samples were made at three distinct sites (repli- cates) in the edge of the gallery forest. At each site, we collected 10 mature inflorescences (with opened flowers) from each plant species along a 50m tran- sect. The inflorescences were carefully put in a plastic bag and cut at the stem base. Once in the laboratory, the spiders from each stem were re- moved, identified to family level, and preserved in 70% ethanol. The spiders were categorized into functional groups based on similarities of foraging behavior. Ambush spiders included only members of the family Thomisidae, whereas hunting spiders included the families Salticidae, Oxyopidae, Clu- bionidae and Anyphaenidae. Web-building spiders and other categories accounted for only 8.2% of all individuals sampled; they were not analysed ac- cording to foraging mode due to small sample sizes, and thus were only included in the analysis of the total number of spiders. The body length between chelicerae and spinnerettes of each individual spider was measured to the nearest 0.1mm. The voucher specimens were deposited in the Colepao Zoologica de Referencia da Universidade Federal de Mato Grosso do Sul (ZUFMS). We compared spider abundance and body size distribution between inflorescences in two pheno- logical stages by collecting stems of Conyza bon- ariensis from the same sampling sites. At each site, we collected 10 young inflorescences which had only flower buds, and 1 0 mature inflorescences with opened flowers. We focussed on C. bonariensis be- cause individual plants of this species with both types of inflorescence were abundant within each sampling site. Sampling procedures were the same as described above. We used one-way ANOVA to compare the abun- dance and mean size of spiders both among the three plant species, and between inflorescences in distinct phenological stages (opened flowers vs. flower buds). We used Tukey’s HSD multiple com- parisons test following ANOVA to determine dif- ferences among plant species. We considered that stems sampled within a site were pseudoreplicates, so the variance associated with them was discarded, resulting in three replicates for each treatment level (see Hurlbert 1984). Our significance level was 0.05. Data were transformed to logio to obtain nor- mality and homogeneity of variances. Influence of plant species on spider abundance and body size distribution, — The total number of spiders on inflorescences differed among the three plant species. Spiders were more abundant on M. latifolia followed by E. hecatanthum, whereas C bonariensis had the lowest number of spiders (Ta- ble 1). The distribution of ambush and hunting spi- ders differed among plant species. Ambush spiders were more common on M. latifolia compared to both C. bonariensis and E. hecatanthum, whereas the abundance of hunting spiders did not differ among plant species (Table 1). Prey availability is regarded as one of the main factors that determine spider abundance (Morse & Fritz 1982; Greenstone 1984; Henschell & Lubin 1997). Although we did not estimate the number of potential prey attracted to the inflorescences of the three plant species, Melanthera latifolia is probably more visited by insects than both C. bonariensis and E. hecatanthum, because it has larger flowers. Inflorescences are regarded as an advertising unit, and several studies showed that larger and more DE SOUZA & MODENA— SPIDERS ON INFLORESCENCES 347 Table 2. — Results of one-way ANOVAs comparing total number of spiders and abundances of ambush and hunting spiders on inflorescences of Conyza bonariensis (Asteraceae) in distinct phenological stages (opened flowers vs flower buds). Spiders Flower buds Opened flowers F P Total spiders 7.3 ± 3.2 4.7 ± 3.1 1.085 0.356 Ambushers 4.0 ± 3.0 2.0 ± 1.0 0.781 0.427 Hunters 2.0 ± 1.7 1.7 ± 2.1 0.172 0.700 opened flowers attract more insects (e.g.. Bell 1985; Cohen & Shmida 1993; Bernays and Chapman 1994; Dafni et al. 1997). Thus, M latifolia may be more attractive to spiders than the other species. Mean body size of spiders sampled on inflores- cences also differed among plant species. Spiders on E. hecatanthum were larger than those on C. bonariensis and M. latifolia (Table 1). Ambush spi- ders on M. latifolia were significantly smaller than those on C. bonariensis and E. hecatanthum. How- ever, hunting spiders found on inflorescences of E. hecatanthum were larger than those on both C bon- ariensis and M latifolia, which sheltered similar sized spiders. Although larger insects may be more frequently attracted by large flowers (Dafni et al. 1997) and could potentially attract larger spiders (Nentwig 1993), studies on vegetative branches show that large spiders are more vulnerable to bird predation (Waldorf 1976; Askenmo et al. 1977). Other structural features of the plants such as branch structural complexity may influence the mi- crohabitat choice by larger spiders due to differenc- es in the availability of refuges against predators (Gunarsson 1990, 1996; Halaj et al. 2000). Thus, there is a need for more studies on the distribution of spiders in distinct inflorescence types, since spi- der groups may respond differently to traits of in- florescences from different plant species. Influence of phenological stage on spider abundance and size distribution.- — The total num- ber of spiders on inflorescences tof C. bonariensis with flower buds was not significantly different from those with opened flowers = 1.085; P = 0.356). The abundances of ambush and hunting spi- ders on both inflorescence types were also similar (Table 2). However, these results should be evalu- ated with caution. Among the plant species studied, the lowest abundance of spiders was recorded on C. bonariensis. The low number of spiders ob- served could obscure differences between inflores- cences in distinct phenological stages, since inflo- rescences of this species were not very attractive for the spiders. Unfortunately, C. bonariensis was the only species in the study area with inflores- cences in both phenological stages, and it was not possible to test this effect on plant species which sheltered larger numbers of spiders. On the other hand, spider body size differed between inflores- cence types. Mean body size of ambush spiders was larger on inflorescences with opened flowers when compared with those bearing flower buds (F,^4 = 17.826; P = 0.013), but no differences in body size were recorded for hunting spiders = 0.009; P = 0.930). These data suggest that, at least for some spider groups, inflorescences in distinct phenologi- cal stages can represent differences in fine-grained qualities of the habitat such as hiding places or prey availability for larger spiders. Several studies of plant-dwelling spiders on veg- etative branches have shown a strong relationship between non-reproductive branch structure and dis- tribution of different spider guilds (Hatley & MacMahon 1980; Scheidler 1990; Uetz et al. 1999; Halaj et al. 2000). Our results suggest that inflores- cence structure and architecture also influences spi- der assemblages, since spiders were more abundant on some types of inflorescence than on others, and mainly because inflorescences of distinct plant spe- cies attracted spiders with different foraging strat- egies. Greco and Kevan (1994) demonstrated that even v/ithout any available prey, Misumena vatia (Clerck) was attracted to yellow color and to a spe- cific plant species, and proposed that these spiders use vision to select microhabitats. In addition, Lou- da (1982) detected differences in the abundances of Peucetia viridans (Hentz) (Oxyopidae) on inflores- cences of two Hapiopappus species (Asteraceae), suggesting that either inflorescence morphology could influence prey availability, or inflorescence type could provide some unknown favorable char- acteristics for those spiders. Inflorescence dwelling spiders can represent an excellent system to clarify questions about which variables influence the dis- tribution of this important arthropod group on the vegetation, because inflorescences have special mi- crohabitat characteristics when compared to non-re- productive branches, potentially influencing the composition and abundance of spider prey and predators attracted to these patches. We are grateful to M. Tanaka for useful com- ments, to the Base de Pesquisas do Pantanal (BEP) for logistic support and CNPq for financial support. The first author had a DCR fellowship from CNPq. 348 THE JOURNAL OF ARACHNOLOGY LITERATURE CITED Askenmo, C., A. von Bromssen, J. Ekman & C. Jansson. 1977. Impact of some wintering birds on spider abundance in spruce. Oikos 28:90-94. Bell, G. 1985. On the function of flowers. Proceed- ings of the Royal Society of London Series B- Biological Sciences 224:223-265. Bernays, E.A. & R.E Chapman. 1994. Host-plant Selection by Phytophagous Insects. Chapman and Hall, New York. Cohen, D. & A. Shmida. 1993. The evolution of flower display and reward. Evolutionary Biology 27:197-243. Dafni, A., M. Lehrer & RG. Kevan. 1997. Spatial flower parameters and insect spatial vision. Bi- ological Reviews 72:239-282. Evans, T.A. 1997. Distribution of social crab spi- ders in eucalypt forests. Australian Journal of Ecology 22:107-111. Greco, C.E & P.G. Kevan. 1994. Contrasting patch choosing by anthophilous ambush predators: vegetation and floral cues for decisions by a crab spider (Misumena vatici) and males and females of an ambush bug (Phymata amehcana). Cana- dian Journal of Zoology 72:1583-1588. Greenstone, M.H. 1984. Determinants of web spi- der species diversity: vegetation structural diver- sity vs. prey availability. Oecologia 62:299-304. Gunnarsson, B. 1990. Vegetation structure and the abundance and size distribution of spruce-living spiders. Journal of Animal Ecology 59:743-752. Gunnarsson, B. 1996. Bird predation and vegetation structure affecting spruce-living arthropods in a temperate forest. Journal of Animal Ecology 65: 389-397. Halaj, J., D.W. Ross & A.R. Moldenke. 1998. Hab- itat structure and prey availabity as predictors of the abundance and community organization of spiders in western Oregon forest canopies. Jour- nal of Arachnology 26:203-220. Halaj, J., D.W. Ross & A.R. Moldenke. 2000. Im- portance of habitat structure to the arthropod food-web in Douglas-fir canopies. Oikos 90: 139-152. Hatley, C.L. & J.A. MacMahon. 1980. Spider com- munity organization: seasonal variation and the role of vegetation architecture. Environmental Entomology 9:632-639. Henschell, J.R., & YD. Lubin. 1997. A test of hab- itat selection at two spatial scales in a sit-and- wait predator: a web spider in the Namib Desert dunes. Journal of Animal Ecology 66:401-413. Hurlbert S.H. 1984. Pseudoreplication and design of ecological field experiments. Ecological Monographs 54:187-21 1. Louda, S.M. 1982. Inflorescence spiders: A cost/ benefit analysis for the host plant, Haplopappiis venetiis Blake (Asteraceae). Oecologia 55:185- 191. Morse, D.H. & R.S. Fritz. 1982. Experimental and observational studies of patch choice at different scales by the crab spider Misumena vatia. Ecol- ogy 63: 172-182. Nentwig, W. 1993. Spiders of Panama. Sandhill Crane Press, Gainesville. Scheidler, M. 1990. Influence of habitat structure and vegetation architecture on spiders. Zoolo- gischer Anzeiger 225:333-340. Uetz, G.W, J. Halaj & A.B. Cady. 1999. Guild structure of spiders in major crops. Journal of Arachnology 27:270-280. Waldorf, E.S. 1976. Spider size, microhabitat selec- tion, and use of food. American Midland Natu- ralist 96:76-87. Manuscript received 24 June 2002, revised 30 June 2003. 2004. The Journal of Arachnology 32:349-352 SHORT COMMUNICATION ITS2 rDNA VARIATION OF TWO BLACK WIDOW SPECIES, LATRODECTUS MACTANS AND LATRODECTUS HESPERUS (ARANEAE, THERIDIIDAE) Daiyuan Zhang', William B. Cook and Norman V. Horner^: Department of Biology: Midwestern State University, Wichita Falls, Texas 76308-2099 USA ABSTRACT. The taxonomic status of two closely related species of Latrodectus, L. mactans and L. Hesperus, has been debated for many years. Based on morphological characteristics and genitalia, some workers consider them as valid species and others as subspecies. This study was conducted to determine whether the internal transcribed spacers 2 (ITS2) of rDNA exhibit sequence differences between the two taxa that could delineate their taxonomic relationship. Individuals of L. mactans and L. Hesperus from six populations were collected and identified based on morphological characteristics. The ITS2 rDNA of nine individuals was sequenced and analyzed. Results indicate that the minimal differences present in the ITS2 sequences are taxonomically insignificant. Keywords^ Theridiidae, Latrodectus, rDNA, internal transcribed spacer 2 The literature reveals conflicting information concerning the taxonomic status of two black wid- ow species, Latrodectus mactans (Fabricius 1775) and L. Hesperus Chamberlin & Ivie 1935 (Araneae, Theridiidae). Taxonomic work based on morpho- logical characteristics has produced controversial conclusions. The two spider taxa were designated as subspecies by Levi (1959), and as separate spe- cies by Kaston (1970, 1978). For this study, the spiders were identified to species following the characteristics outlined by Kaston. Specimens of each species are on deposit in the Invertebrate Col- lection at Midwestern State University, Wichita Falls, Texas. The rRNA genes have long been recognized as attractive markers for phylogenetic studies (Hillis & Dixon 1991). These genes are organized in clus- ters of repeated units, each of which consists of coding sequences, and several transcribed and non- transcribed spacer regions (NTS). The transcription units include the 18S, 5.8S, and 28S genes as well as external and internal transcribed spacers (ETS and ITS). Coding regions and spacers differ greatly in their rate of evolution, and hence the rDNA clus- ters have the potential to reveal phylogenetic rela- ' Current Address: Dept, of Biological Science, University of North Texas, Denton, Texas 76203- 5220. ^ Corresponding author. tionships at many taxonomic levels. The level of divergence observed in the spacer regions is appro- priate for detecting differences between specific in- dividuals, which provides a potentially useful mark- er with which to study the relationships of populations and closely related species (Cerbah & Souza 1998; Hedin 1997; Vogler 1994). For ex- ample, Vogler (1994) separated tiger beetles (Cicin- dela dorsalis)', Hedin (1997) separated cave spider populations and species (Nesticus)', Harris & Cran- dall (2000) separated closely related species and populations of freshwater crayfishes (Decapoda, Cambaridae). This preliminary study of ITS2 rDNA variation was conducted using individuals from three popu- lations for each species. Latrodectus mactans indi- viduals were collected in Texas from Wichita (34°00'N, 98°42'W), Brown (3U45'N, 99°00'W) and Kimble (30°29'N, 99°46'W) Counties. Latro- dectus Hesperus individuals were collected from Eddy County, New Mexico (32°52'N, 104°45'W), Brewster (29°33'N, 103°47'W) and Garza (33°10'N, 101°20'W) Counties of Texas. Genomic DNA was isolated from leg IV of each individual spider using QIAGEN DNAeasy Tissue Kit (Qiagen, Inc., Va- lencia, CA). The DNA concentration was measured using a UV spectrophotometer (Pharmacia, Ultra- spec III). The ITS2 region of rDNA was amplified using 5.8S primer (5'-GGGACGATGAAGAACGCAGC- 349 350 THE JOURNAL OF ARACHNOLOGY Lat CAGCTGCGAGACTTGGTGTGAATTGCAGGACACATTGAGCACTGATTTTTCGAACGCGCA Eno CAGCTGCGAGACT-GGTGTGAATTGCAGGACACATTGAGCACTGATTTTTCGAACGCGCA ************* ************■*■***•*■****■*•■*•***■*•***■*•******•*■****■*■■*** Lat TTGCGCCCTCT-~~--TCCCCGGGGGCTCGCCTGTCTGAGGGTCGGATAAGACTTACCGA Eno TTGCGGCCTCGGGTCCTGCCCGGGGCCTCGCCTGTCTGAGGGTCGGATAAGACTTGCAAA ***** **** * ******* ***************************** * * Lat GGAGGGTTTCCCTCCCA-TTGGCTGCATCGGATGG---TGATCATCCGTCTGCCTAAGGT Eno GGAAAGTTCGCTTTCCACTTGGCCGAACCGGATCGCTTTTGTGGTTCGCCGGCTTAAGGT *** *** * * *** ***** * * ***** * * * ** * ** ****** Lat CTTCGTCCGCGAATCCCCCTGGCCGAGAAGCGTCGCTCC--~TCGAATAGGTCCAGCATA Eno TTACG— — GATCCTCCCCGGCCGAGAAGCGTGGCTCCCACTTGACCAACGCTCGCGG- * ** ** * *** ************* ***** * ** * * ** Lat GAGGGAGAACGACTGA-TCGTCTCCCCGCCGACTGGAGCTGGAGCTGATGACGCAGAGAG Eno GCTGGAGAA-GACTGAGTAGTTTCTCCGCTGAAGCGAGCGACGGGAGCGAGCGCAGAGAG * ****** ****** * ** ** **** ** **** * * ********* Lat TGTGCCGGGAAGAAGGGATGGTCGGATGGTCCGCTTGGGTGCTTTCGTGCTAATGCGTCC Eno CTTGCCGAGGAGA— GATCCTCCAC-- — — — — — — — — GCGTGC ***** * *** *** *** * **** * Lat CTCGTTCGTATAACGGCGTTGAACTACGTCCTGAATTGAGGGTTCGCAGCGAAAGGTCAC Eno C------ — ---AC — — -TGGCACGGTAAACTCAA * ** * *** * ** *** Lat GACTGATATTCATATCTGTCGACCTCAGATCAGACGAGATGACCCGCTGAATTTAA Eno AA— -- — ATAT-TGTCGACCTCAGATCAGACGAGATGACCCGCTGAATTTAA * **** **************************************** Figure 1. — Alignment of ITS2 regions from Latrodectus spp. and Enoplognatha ovata. Lat. = consensus sequence of eight Latrodectus mactans and Latrodectus hesperus individuals. Eno. = sequence of L. ovata (Araneae, Theridiidae). Underlined ITS2 sequences are flanked by 5.8S and 28S sequences. Asterisks indicate identical nucleotide positions. Alignment generated by CLUSTALW 1.4. 3') and 28S primer (5'-TCCTCCGCTTATTGA- TATGC-3') (Hillis et al. 1996). Two nanograms of spider genomic DNA were used for each reaction and PCR amplification was performed in 100 piL, using the following profile: 5 min at 94 °C, 40 cy- cles (1 min at 94 °C, 2 min at 45 °C, 1.5 min at 72 °C), and 7 min at 72 °C. The products were assessed by mini-gel electrophoresis using 5 |jlL aliquots. Successful amplifications generated a single 0.5 kb product. After purification, the PCR products were digested with LcoK I and Hind III and ligated into pUC19 vector. The recombinant plasmids were transformed into L. coU DH5a cells. Sequences of both strands were determined by Northwoods DNA, Inc. (Center for Research and Innovation, 44526 CNTY Rd 3, Becida, MN 56678). Results were re- ceived as unprocessed nucleotide sequences and an- alyzed manually. Some visual adjustments were made. The individual sequences were aligned using CLUSTAL W 1.4 (Higgins & Sharp 1988) and con- sensus sequences were obtained. First, the extent of the intergenic region between the 5.8S and 28S coding regions referred to as ITS2 was determined by similarity with the published se- quences of Lnoplognatha ovata (Araneae; Theridi- idae) (Fig. 1) (Tan & Gillespie 1999). The length of the ITS2 region in different clones of L. mactans and L. hesperus is —360 bp. Three individuals (LMWl, LMW2, LMW3) of the LMW population (L. mactans from Wichita County, Texas) were se- lected to analyze the variation in one population. For individual LMW3, four separate clones were sequenced to test variation within one individual. For the other populations, one individual from each was randomly selected for PCR amplification and one or more independent clones covering the ITS2 were sequenced from each of these individuals. Five variable nucleotide positions were found in ZHANG ET AL.— VARIATION IN LATRODECTUS 351 1 2 3 4 LMWl C T T T LMW2 c T T C LMJ T T A T LMB T A T T LHM T A T T LMW3 T T A T LHD T T T T LHX T T A T Figure 2. — Alignment of Variable ITS2 sites from eight Latrodectus individuals. Variable nucle- otides occur at four positions within the Latrodectus consensus sequences in Figure 1:1 = 124; 2 = 153; 3 = 227; 4 = 422. LMW = L. mactans, Wichita County, Texas; LMJ = L. mactans. Llano County, Texas; LMB = L. mactans. Brown County, Texas; LHM = L. hesperus, Eddy County, New Mexico; LHD = L. hesperus, Brewster County, Texas; LHX = L. hesperus, Garza County, Texas. the alignment of sequences from individual LMW3. Three variable nucleotide positions were found in the alignment of three individuals from Wichita County population of L. mactans. Three variable nucleotide positions were found in the alignment of three L. mactans populations. Only one variable nu- cleotide position was found among three popula- tions of L. hesperus. The consensus sequences of eight individuals from six populations varied at four nucleotide positions (Fig. 2). The phylogenetic analysis of the aligned se- quences was performed as described by Templeton et al. (1992) using the TCS software package (Clement et al. 2000). ITS2 sequences from eight Latrodectus individuals collapsed into five haplo- types (Fig. 3). The three LMW individuals repre- sented three different haplotypes on two different branches from a L. hesperus node. LMW3 and two L. mactans individuals from outside Wichita Coun- ty represented a single haplotype. Latrodectus hes- perus sequences collapsed into two haplotypes, one of which supported the two L. mactans branches. Even though all the clones, individuals, popula- tions and two taxa showed some level of sequence variation in the ITS2 region of rDNA, the separa- tion between species was not well supported on these grounds. For L. mactans, variation within an individual is 1.4%, variation among individuals and that among populations are each 0.83%. The L. hes- perus populations exhibit only 0.27% variation. The variation between two taxa is 0.83% and does not distinguish these two species. The same result was found in the ITS2 research of mosquitoes. Wesson et al. (1992) reported 0.46% variation within 10 Figure 3. — Phylogenetic network of five haplo- types from ITS2 sequences of eight Latrodectus in- dividuals. Open shapes, representing the haplo- types, indicate the included Latrodectus individuals. Analysis was performed by TCS (Clements et al. 2001). clones of ITS2 from a single mosquito, Aedes simp- soni, while intraspecific variation in A. aegypti was only 1.17%. The differentiation even within a single individual may be caused by the existence of poly- morphisms among repeat units of rDNA. In a phylogenetic analysis of ITS2 sequence var- iations, L. mactans individuals from Wichita Coun- ty represented as many haplotypes as the other five individuals, which included representatives of both L. hesperus and L. mactans. Furthermore, the three LMW haplotypes represented two distinct branches from LHM. Assuming that the Wichita County L. mactans individuals reflect a typical level of re- gional variation, ITS2 sequences do not offer a re- liable means of distinguishing between L. mactans and L. hesperus populations. Only those characters that are diagnostic for all individuals of an entire population can be used in the phylogenetic reconstruction of populations (Vo- gler 1994). Further exploration of the molecular taxonomy of these taxa will require additional data, including sequence comparisons of mtDNA, ITSl DNA, plus other nuclear genes. LITERATURE CITED Cerbah, M. & T. Souza. 1998. Molecular phylogeny of the genus Hypochaeris using internal tran- scribed spacers of nuclear rDNA: Inference for chromosomal evolution. Molecular Biology and Evolution 17:284-291. Clement, M., D. Posada & K.A. Crandall. 2000. TCS: a computer program to estimate gene ge- nealogies. Molecular Ecology 9:1657-1659. Harris, D.J. & K.A. Crandall. 2000. Intragenomic 352 THE JOURNAL OF ARACHNOLOGY variation within ITSl and ITS2 of freshwater crayfishes (Decapoda: Cambaridae): Implications for phylogenetic and microsatellite studies. Mo- lecular Biology and Evolution 17:284-291. Hedin, M.C. 1997. Molecular phylogenetics at the population/species interface in cave spiders of the Southern Appalachians (Araneae: Nesticidae: Nesticus). Molecular Biology and Evolution 14: 309-325. Higgins, D.G. & RM. Sharp. 1988. CLUSTAL: A package for performing multiple sequence align- ment on a microcomputer. Gene 73:237-244. Hillis, D.M. & M.T. Dixon. 1991. Ribosomal DNA: molecular evolution and phylogenetic inference. The Quarterly Review of Biology. 66(4):411- 453. Hillis, D., M.C. Moritz & B.K. Mable. 1996. Mo- lecular Systematics. Sinauer Associates, Inc. Sunderland, Massachusetts. Kaston, B.J. 1970. Comparative Biology of Amer- ican Black Widow Spiders. Transactions of San Diego Society of Natural History. 16:33-32. Kaston, B.J. 1978. How to Know the Spiders. 3*^^* ed. Wm. C. Brown Company Publishers, Du- buque. Levi, H.W. 1959. The spider genus Latrodectus (Araneae: Theridiidae). Transactions American Microscopical Society. 78(l):7-43. Tan, A.M. & R.G. Gillespie. 1999. Paraphyly of the Enoplognatha ovata group (Araneae, Theridi- idae) based on DNA Sequences. Journal of Ar- achnology 27:481-488. Templeton, A.R., K.A. Crandall & C.E Sing. 1992. A cladistic analysis of phenotypic associations with haplotypes inferred from restriction endo- nuclease mapping and DNA sequence data. III. Cladogram estimation. Genetics 132:619-633. Vogler, D. 1994. Evolution and phylogenetic infor- mation content of the ITS-1 region in the tiger beetle Cicindela dorsalis. Molecular Biology and Evolution 11:393-405. Wesson, D.M., C.H. Porter & EH. Collins. 1992. Sequence and secondary structure comparisons of ITS rDNA in mosquitoes (Diptera: Culicidae). Molecular Phylogenetics and Evolution 1:253- 269. Manuscript received 23 September 2002, revised 11 August 2003. INSTRUCTIONS TO AUTHORS (revised October 2003) General: Manuscripts are accepted in English only. Authors whose primary language is not English may consult the editors for assistance in obtaining help with manuscript preparation. 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Figures should be arranged so that they fit (vertically and horizontally) the printed journal page, either one column or two columns, with a minimum of wasted space. When reductions are to be made by the printer, pay particular attention to width of lines and size of lettering in line drawings. Multiple photos assembled on a single plate should be mount- ed with only a minimum of space separating them. In the case of multiple illustrations mounted together, each illustration must be numbered sequentially rather than given an alphabetic sequence. Written on the back should be the name(s) of author(s) and an indication of top edge. Indicate whether the illustration should be one column or two columns in width. The overall dimensions should be no more than 1 1 inches (28 cm) X 14 inches (36 cm). Larger drawings present greater difficul- ty in shipping and greater risks of damage for which the Journal of Arachnology assumes no responsibility. In manu- scripts for review, photocopies should be included, and should be reduced to the exact measurements that the author wants to appear in the final publication. Make notations in the text mar- gins to indicate the preferred position of illustrations in the printed text. Color plates can be printed, but the author must assume the full cost, currently about $600 per color plate. Legends for illustrations should be placed together on the same page(s) and separate from the illustrations. Each plate must have only one legend, as indicated below: Figures 1-4. — A-us x-us, male from Timbuktu: 1. Left leg; 2. Right chelicera; 3. Dorsal aspect of genitalia; 4. Ventral aspect of abdomen. Figures 27-34. — Right chelicerae of species of A-us from Timbuktu: 27, 29, 31, 33. Dorsal views; 28, 30, 32, 34. Prolateral views of moveable finger; 27, 28. A-us x-us, holo- type male; 33, 34. A-usy-us, male. Scale = 1.0 mm. Assemble manuscript for mailing. — Assemble the sepa- rate sections or pages in the following sequence; title page, abstract, text, footnotes, tables with legends, figure legends, figures. Page charges, proofs and reprints. — Page charges are vol- untary, but non-members of AAS are strongly encouraged to pay in full or in part for their article ($75/journal page). The author will be charged for changes made m the proof pages. Reprints are available only from the Allen Press and should be ordered when the author receives the proof pages. Allen Press will not accept reprint orders after the paper is published. The Journal of Arachnology also is publishea by BioOne. Therefore, you can download the PDF version of your article from the BioOne site or the AAS site if you are a member of AAS or if your institute is a member of BioOne. PDF’s of arti- cles older than one year will be freely available from the AAS website. SHORT COMMUNICATIONS Short Communications are usually limited in length to three journal pages, including tables and figures. They will be print- ed in a smaller (10 point) typeface. The format for these is less constrained than for feature articles: the text must still have a logical flow, but formal headings are omitted and other deviations from standard article format can be permitted when warranted by the material being covered. CONTENTS The Journal of Arachnology Volume 32 Featured Articles Number 2 Life histories of four speeies of scorpion in three families (Buthidae, Diplocentridae, Vaejovidae) from Arizona and New Mexico by Christopher A. Brown 193 Population dynamics of an isolated population of the harvestman Ilhaia cuspidata (Opiliones, Gonyleptidae), in Araucaria Forest (Curitiba, Parana, Brazil) by Luiz Augusto Macedo Mestre & Ricardo Pinto-da-Rocha 208 Phenology of linyphiids in an old-growth deciduous forest in central Alberta, Canada by Christopher M. Buddie & Michael L. Draney 221 A revision of the spider genus Calileptoneta Platnick (Araneae, Leptonetidae), with notes on morphology, natural history and biogeography by Joel M. Ledford 231 Three new species of the spider genus Phrurolithus from China (Araneae, Corinnidae) by Chang-Min Yin, Darrell Ubick, You-Hui Bao & Xiang Xu 270 The movement and activity patterns of similar-sized adult and juvenile crab spiders Misumena vatia (Araneae, Thomisidae) by Heather L. Sullivan & Douglass H. Morse 276 The systematics of the Eremobates scaber species-group (Solifugae, Eremobatidae) by Jack O. Brookhart & Paula E. Cushing 284 Assemblages of spiders on models of semi-arid shrubs by Michael W. Heikkinen & Janies A. MacMahon 313 Group size does not influence growth in the theraphosid spider Hysterocrates gigas (Araneae, Thersphosidae, Eumenophorninae) by Melissa M. Varrecchia, Vanessa A. Gorley, & Samuel D. Marshall . . 324 Short Communications A new species and a new synonymy in the spiny orb-weaver spider genus Micrathena (Araneae, Araneidae) by Marcelo O. Gonzaga & Adalberto J. Santos 332 Color dimorphism in adults and juveniles of Buitinga safura (Araneae, Pholcidae) by Bernhard A. Huber & Sascha Hopf 336 Pardosa milvina (Araneae, Lycosidae) spiderling movement in the presence of conspecific and heterospecific silk and excreta by Matthew H. Persons & Erin C. Lynam 341 Distribution of spiders on different types of inflorescences in the Brazilian pantanal by Andrea Lucia Teixeira de Souza & Erica de Souza Modena 345 ITS2 rDNA variation of two black widow species, Latrodectus mactans and Latrodectus hesperus (Araneae, Theridiidae) by Daiyuan Zhang, William B. Cook & Norman V. Horner 349 USERNAME: norman04 PASSWORD: spider04