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Arachnida, Opiliones) 1998. The Journal of Arachnology 26:296-302 LEG AUTOTOMY AND ITS POTENTIAL FITNESS COSTS FOR TWO SPECIES OF HARVESTMEN (ARACHNIDA, OPILIONES) Cary Guffey' : Department of Biology, PO Box 42451, University of Southwestern Louisiana, Lafayette, Louisiana 70504 USA ABSTRACT . Leg autotomy often confers immediate benefits on the animal losing its legs, such as escape from a predator, while costs are usually less obvious and accrue long after the leg is lost . I conducted a survey to determine the prevalence and characteristics of leg autotomy in two species of harvestmen, Leiobunum nigripes Weed 1892 and L. vitlalum Say 1821, from May-December 1996 at Chicot State Park, Evangeline Parish, Louisiana . Nearly half of all individuals found were missing at least one leg . There was no significant difference in the median number of legs between months for L. nigripes, but differences were found among several months for L. vittatum. Either of the second legs was most likely to be lost in both species . These results indicate that leg autotomy is common in harvestmen . Furthermore, these results suggest that the second legs are not as crucial to the survival of harvestmen as previously believed. Leg autotomy may result in a reduction of potential fitness for individuals, but harvestmen may choose to incur these costs rather than risk a catastrophic loss of fitness (e .g ., death) ; that is, leg autotomy may be a bet-hedging strategy for harvestmen . Autotomy of appendages is widely known therefore, there are no costs associated with for a variety of animal groups, including mol- regeneration, such as allocation of nutrients to luscs (Edmunds 1966), crustaceans (Juanes & new tissue . However, harvestmen also are not Smith 1995), arachnids (e .g ., spiders : Forma- able to recover the benefits that they originally nowicz 1990 ; harvestmen : Kaestner 1968), in- had with the full complement of eight legs . sects (Carlberg 1994), echinoderms (e.g ., Thy- Because harvestmen do not regenerate their one briareus LeSueur 1824 : Smith & legs, when confronted with predators they Greenberg 1973), salamanders (Wake & Dres- should be under natural selection to weigh the ner 1967), and lizards (Arnold 1984) . The ef- benefits of leg autotomy against the future fect that autotomy has on the fitness of an in- costs associated with autotomy, such as sub- dividual depends on the sum of the benefits sequent encounters with predators, loss of mo- and costs of appendage loss . Benefits may be bility, loss of foraging ability, or loss of mat- immediate or nearly so and include distraction ing opportunities . of predators (Arnold 1984), escape from a The second pair of legs is the longest in predator's grasp (Arnold 1984), and escape Leiobunum (Kaestner 1968 ; Edgar 1990), and from traps (e .g ., spider webs) . Costs are these legs are believed to contain the main spread over a longer period of time and in- sensory organs of these animals (Comstock clude loss of mobility or balance, reduced 1920 ; Edgar 1963 ; Cloudsley-Thompson ability to escape subsequent encounters with 1968) . In one study, harvestmen that had lost predators, decrease in social status, divergence the second pair of legs were reported to be of energy resources to replacement of the lost more reluctant to move, eat, drink, or mate appendages, decrease in mating success, and (Sankey & Savory 1974) . Cloudsley-Thomp- even death in some instances (reviewed in Ed- son (1968) wrote that the loss of both of the munds 1974 ; Arnold 1984, 1988 ; Juanes & second pair of legs quickly results in death . Smith 1995) . Given the reported importance of the second Harvestmen autotomize their legs but do pair of legs, I predict that harvestmen should not regenerate them as either juveniles or be particularly reluctant to autotomize this adults (Comstock 1920 ; Kaestner 1968) ; pair compared to the other pairs . In general, three different life histories can ' Current address : Our Lady of the Lake University, be found in Opiliones (Todd 1949) . (1) Eggs 411 SW 24th St ., San Antonio, Texas 78207 USA . laid the previous autumn hatch in the spring . 296 GUFFEY--LEG AUTOTOMY IN HARVESTMEN 297 The animals mature over the summer and lay corded location, number of legs, which legs eggs in the autumn, then adults may or may were missing, and whether or not collected in- not die at the first frost (e.g., Leiobunum ni- dividuals were found in the copulatory posi- gripes Weed 1892) . This results in an overlap tion. Both species in this study are sexually of generations in some species (e.g., Phalan- dimorphic, but the characters used to identify gium opilio Linnaeus 1758 : Clingenpeel & the sexes of L. nigripes (size, color; pers. obs.) Edgar 1966) . (2) Eggs hatch in the autumn are not as reliable as that used for L. vittatum and the young overwinter . The young mature (size of pedipalps ; Davis 1935) . Any animals during the following summer, lay their eggs, for which the sex was uncertain were removed then die before the clutches hatch . Clingen- from comparisons based on sex . I released all peel & Edgar (1966) used populations of animals near the sites at which they were col- Leiobunum politum Weed 1889 and L. vitta- lected. A previous mark-recapture study in- tum Say 1821 from Michigan as examples, but dicated that <10% of marked animals were the latter species appears to follow pattern I caught at the site at which they were marked in south central Louisiana (pers . obs .). (3) As after 24 h and < I % were recaptured after two in the previous case, the eggs hatch in the au- weeks (unpubl . data); therefore, pseudorepli- tumn and the young overwinter. Eggs are laid cation due to resampling should have been the following autumn, but the adults do not negligible. die until after the eggs hatch, resulting in an Comparisons between months .-I used a overlap of generations (e .g., L. townsendi Kruskal-Wallis one-way ANOVA by rank Weed 1893 : Cokendolpher et al . 1993). Har- (Siegel & Castellan 1988) to determine if vestmen are ametabolous, and the young un- there was any significant difference for each dergo 5-8 molts before reaching adulthood species in the number of legs present among (Edgar 1971 ; Kaestner 1968) with the first months . When a significant difference was de- molt taking place within hours of hatching tected, I used a multiple comparisons test (Edgar 1971) . (Siegel & Castellan 1988) to locate significant Both L. vittatuin and L. nigripes were col- differences between months . These tests were lected throughout the year at Chicot State two-tailed and the test statistics were adjusted Park, Evangeline Parish, Louisiana ; but it was for ties with a = 0 .05 . common to find only adult L. vittalum from Comparisons of leg pairs .-I summed the late November to early January and to find number of times that either leg of a particular only juvenile L. nigripes from early January pair was autotomized and compared these ob- to mid-February. Leiobunum nigripes served values to an expected value obtained emerged 1--2 months earlier than L. vittatum. by assuming that each pair was equally likely In this study, I conducted field observations to lose a member. The probability (P) that ei- of L. nigripes and L. vittatum to determine the ther leg of any particular pair would be lost prevalence of leg autotomy in these species, was 0.25. This value was calculated as follows which legs are most likely to be autotomized, P(X,, or Xk) = P(X,) + P(X1,) _ % + /K = /K and what costs might be associated with leg = 0.25 autotomy. 1 predicted that : (1) the proportion of individuals missing legs should increase where X is any particular leg pair, L indicates with age, (2) the second legs should be the left, and R indicates right . This null hypothesis least likely to be autotomized, and (3) indi- was evaluated for both species with a Chi- viduals found mating should be more likely to square goodness of fit test (Freund & Simon have all eight legs than those individuals 1992) and compared to a = 0 .05. found alone . I compared the number of individuals miss- ing both legs of the second pair to an expected METHODS probability of 0 .036 (again assuming an equal Study animals.-I observed juveniles and likelihood that any leg would be autotomized) . adults of L. nigripes and L. vittatum at Chicot This probability was calculated as follows State Park, Evangeline Parish, Louisiana be- P(X) = P(X,) and P(X,) = P(X,)P(X2) _ tween May and December 1996 . Individuals (/) ('/7) = 0.036 were collected from cabins, vegetation, and 10.2 X 10.2 X 50.0 cm wooden posts . I re- where X is any particular leg pair. I used a 298 THE JOURNAL OF ARACHNOLOGY 0 .6 Leiobunum nigripes 0 .7 427 Males Females L . nigripes L . vittatum 167 0 .6 ze 0 .5 0 0 .5 0 61 0 .4 0 0 .3 0 .4 0 w 0 .2 s 7 0 0 .i 2 2 0 0 .3 223 0 .0 - a e4 4 5 6 7 8 4 5 6 7 8 0 2 Number of legs present a Leiobunum vittatum 0 .2 0 .7 43 71 0 .6 4 0 .5 0 .1 0q 17 „ 0 .4 0 131 92 0 0 .3 0 .0 W '' 0 .2 4 5 6 7 8 40 Number of legs present 0 .1 I z o Figure l .-The incidence of leg autotomy in two 0 .0 4 5 6 7 8 4 5 6 7 8 species of harvestmen from Evangeline Parish, Number of legs present Louisiana, May-December 1996 . The proportion of individuals with a given number of legs was not Figure 2 .-A comparison of leg number by spe- significantly different between the two species .
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