The in flu ence of trap type on eval u at ing pop u la tion struc ture of the semifossorial and so cial ro dent Octodon degus

Acta Theriologica 54 (4): 311 32, 29. PL ISSN 1-751 DOI: 1.498/j.at.1-751.47.28 The in flu ence of trap type on eval u at ing pop u la tion struc ture of the semifossorial and so cial ro dent Octodon degus Jo seph R. BURGER, Adrian S. CHESH*, Rodrigo A. CASTRO, Liliana ORTIZ TOLHUYSEN, Ignasi TORRE, Luis A. EBENSPERGER and Loren D. HAYES Bur ger J. R., Chesh A. S., Cas tro R. A., Ortiz Tolhuysen L., Torre I., Ebensperger L. A. and Hayes L. D. 29. The in flu ence of trap type on eval u at ing pop u - la tion struc ture of the semifossorial and so cial ro dent Octodon degus. Acta Theriologica 54: 311 32. Trap type may in flu ence cap tures of in di vid u als in dif fer ent age-sex cat e go ries in small mam mal stud ies, re sult ing in bi ased pop u la tion and de mo graphic in for ma tion. We de ployed 4 live trap types at bur row sys tems of the ro dent, Octodon degus Molina, 1782, in cen tral Chile to de ter mine trap ef fi cacy in cap tur ing in di vid u als of 6 de mo graphic cat e go ries. We cap tured 2672 in di vid u als in 17 79 trap days (15.1% trap ping suc cess). Tom a hawks were the most ef fi cient trap cap tur ing half of in di vid u als dur ing both years, fol lowed by mesh traps, large traps, and me dium traps in 25. All trap types equally sam pled sexes. and me dium traps pro vided sim i lar de mo graphic struc ture, where half of the in di vid u als cap tured were pups; Tom a hawk traps sam pled more adults than pups. Rel a tive cap tures of pups were sim i lar across dif fer ent trap types, sug gest ing that pups are equally sam pled by each of the de ployed trap types. Rel a tive cap tures of adults were lower in traps, sug gest ing that this age class avoided solid-walled traps. For Octodon degus, the sole use of Tom a hawk traps may pro duce suf fi cient, un bi ased de mo graphic data. Only 4 trap mor tal i ties oc curred (.15%). Re search ers may min i mize trap mor tal ity with out com pro mis ing suf fi cient de mo graphic sam pling by trap ping dur ing peak an i mal ac tiv ity. De part ment of Bi ol ogy, Uni ver sity of Lou i si ana at Mon roe, 7 Uni ver sity Av e nue, Mon roe, LA 7129, USA, e-mail: robbieburger@mac.com (JRB, ASC, LDH); Departamento de Ecología, Facultad de Ciencias Biológicas & Centro de Estudios Avanzados en Ecología y Biodiversidad, Pontificia Universidad Católica de Chile, San ti ago, Chile 6513677 (RAC, LOT, LAE); Museu de Granollers-Ci ncies Nat u rals, 842 Granollers, Spain (IT) Key words: de mog ra phy, live-trap ping, small mam mals, trap, sociality, Tom a hawk trap In tro duc tion Long-term de mo graphic stud ies of small mammals are im por tant for un der stand ing pop u la - tion dy nam ics and be hav ior (Metcalf and Pavard 27). The suc cess of these stud ies de pends on the re search ers abil i ties to cap ture in di vid u als rep re sent ing all de mo graphic groups pres ent in * Pres ent ad dress: De part ment of Zo ol ogy, Mi ami Uni ver sity, Ox ford, OH 4556, USA [311]

312 J. R. Burger et al. the pop u la tion. Un equal sam pling of de mo g- raphic groups could re sult in er ro ne ous es ti mates of den sity, sur vival, sex ra tios, and adult-tooff spring ra tios. Ad di tion ally, bi ased sam pling in a so cial spe cies can limit re search ers abil i ties to de ter mine so cial group size and com po si tion. Eval u at ing the ef fec tive ness of mul ti ple trap types in cap tur ing in di vid u als from dif fer ent sex and age classes in a pop u la tion is crit i cal, yet few stud ies have ad dressed this is sue. Many fac tors can in flu ence small mam mal trap suc cess, in clud ing those that re search ers can con trol such as trap type (Wiener and Smith 1972, Slade et al. 1993, O Farrell et al. 1994). Me chan i cal dif fer ences in live traps, such as trea dle sen si tiv ity, can in flu ence trap suc cess among age-size de mo graphic groups (Wiener and Smith 1972, Rose et al. 1977, Slade et al. 1993). Ad di tion ally, a num ber of un con trol la ble fac tors in trin sic to the study or gan ism may in - flu ence trappability such as be hav ior (ie, trap shy vs trap prone in di vid u als), age class, and body mass of in di vid u als (Neal and Cock 1969, Gliwicz 197, An thony et al. 25). These fac tors are par tic u larly chal leng ing to re search ers be - cause vari abil ity in trap ef fi cacy may in tro duce bi ases (Boonstra and Rodd 1982, Maly and Cranford 1985). There fore, a com bi na tion of mul ti ple trap types might be nec es sary to ac cu - rately sam ple small mam mals of vari able sizes (Szaro et al. 1988, O Farrell et al. 1994, Iriarte et al. 1989, An thony et al. 25, Santos-Filho et al. 26). Two com monly used types of live traps in small mam mal stud ies are the (H. B. Inc., Tal la has see, FL) and Tom a hawk traps (Tom a hawk Live Trap Co., Tom a hawk, WI). Al though sev eral stud ies have com pared the trap ef fi cacy of live traps to other trap types in cap tur ing small mam mals (Sealander and James 1958, Slade et al. 1993, O Farrell et al. 1994, An thony et al. 25), few stud ies have com pared the rel a tive ef fec tive ness of vs Tom a hawk live traps. Santos-Filho et al. (26) found Tom a hawk traps to be es sen tial in sam pling a small mam mal com mu nity in the Neotropics, cap tur ing 6 spe cies that were com - pletely missed us ing a suite of, snap, and pit fall traps. Iriarte et al. (1989) re ported dif fer ences in me dium live traps vs Tom a hawk live traps in a long-term com mu nity study in cen tral Chile. Tom a hawk traps were poor at sam pling small ro dents and mar su pi als and pro vided lower pop u la tion den sity es ti - mates for larger spe cies, in clud ing Octodon degus Molina, 1782. No study, to our knowl edge, has com pared the trap efficacies of mul ti ple trap types, in clud ing s and Tom a hawks, in sam pling in di vid u als of a sin gle spe cies to de ter - mine so cial group and pop u la tion de mog ra phy. Dif fer ences in trap de sign may in flu ence trappability and trap suc cess. traps dif fer greatly from solid-walled traps in the amount of light al lowed into the trap. The in ter nal, dark na ture of solid-walled traps may re - sem ble a bur row or other ref uge. In con trast, mesh traps (eg Tom a hawk) al low vis i bil ity of the ex ter nal en vi ron ment. In a study in volv ing the noc tur nal and sol i tary dusky footed woodrat Neotoma fuscipes, Laudenslayer and Fargo (22) had more cap tures us ing solid-walled traps than with mesh Tom a hawks when trap - ping at woodrat nest sites, but the op po site was true when trap ping on a grid. O Farrell et al. (1994) found mesh traps to be more ef fec tive than solid-walled traps in cap tur ing a num ber of small mam mal spe cies in a va ri ety of hab i tats and geo graph ical lo ca tions. To date, no study has de ter mined the ef fec tive ness of mul ti ple trap types, in clud ing mesh and solid-walled traps in sam pling in di vid u als of dif fer ent de mo graphic cat e go ries within a spe cies. The suc cess of dif fer ent trap types may also de pend on the be hav ior of the study or gan ism. For ex am ple, semifossorial ro dents spend a con - sid er able amount of time un der ground, emerg - ing from bur rows to for age. Cap tur ing these an i mals may be dif fi cult, and bur row counts may re sult in bi ased pop u la tion den sity es ti - mates (Powell et al. 1994, Van Horne et al. 1997). Trap place ment may in flu ence cap tures in mam mals that are microhabitat spe cial ists (Knowles and Bur ger 28). For ex am ple, Loeb et al. (1999) found that trap place ment in trees

Trapping population structure of Octodon degus 313 was nec es sary to doc u ment ar bo real spe cies. In con trast, trap place ment at bur row open ings or along run ways may be nec es sary to ef fi ciently cap ture semifossorial spe cies. There fore, knowl - edge of trap ef fec tive ness for cap tur ing bur row - ing ro dents is es pe cially im por tant for eco log i cal stud ies. Degus Octodon degus are me dium-sized (adults ~17 3g), caviomorph ro dents en - demic to Chile (Woods and Boraker 1975). Degus are di ur nally ac tive and semifossorial, emerg ing from bur rows at day break to for age (Kenagy et al. 22, Ebensperger et al. 24, Hayes et al. 27). Ac tiv ity pat terns are bi modal dur ing the aus tral spring and unimodal dur ing the aus tral win ter (Kenagy et al. 22). Degus are so cial, liv ing in groups con sist ing of sev eral males and re pro duc tive fe males, which fre quently use the same bur row sys tems (Ebensperger et al. 24). These bur row sys tems are ex ten sive, in volv ing nu mer ous bur row open ings (Hayes et al. 27, Hayes et al., in press, Ebensperger et al. 29). Thus, ef fi cient trap ping meth ods are re quired to quan tify the size and com po si tion of so cial groups. Not sur pris ingly, pre vi ous stud ies in - volv ing degus in di cated that trap type might in - flu ence cap ture suc cess. Fulk (1976) and Jaksic et al. (1981) ques tioned the ad e quacy of me dium traps in sam pling larger degus. On the other hand, Iriarte et al. (1989) stated that young degus might be able to es cape from mesh Tom a hawk traps, re sult ing in bi ased den sity es - ti mates. There fore, it may be nec es sary to de - ploy mul ti ple trap types to ac cu rately sam ple de mog ra phy of so cial groups and pop u la tions of this spe cies. As part of an on go ing study of degu sociality and space use (Hayes et al. 27, Hayes et al., in press, Ebensperger et al. 29), we de ployed 4 trap types to de ter mine the ef fi cacy of to tal cap - tures and cap tures of in di vid u als from 6 demographic categories. Our objective was to determine what type of trap, or com bi na tion of traps, pro - duced an ac cu rate rep re sen ta tion of so cial group and pop u la tion struc ture. We spe cif i cally tested the null hy poth e sis that in di vid u als from all de - mo graphic cat e go ries and to tal cap tures were equally likely to be cap tured by each trap type used. Ma te rial and meth ods Study area Our study took place in cen tral Chile at the Estación Ex per i men tal Rinconada de Maipú (33 23 S, 7 31 W, al ti - tude = 495 m), a field sta tion of the Universidad de Chile. Med i ter ra nean-type cli mate with cool wet win ters and warm dry sum mers char ac ter izes the site. The hab i tat is Chil ean matorral fea tur ing an open land scape with scat tered shrubs and a grass and forb her ba ceous com po si tion. Dur ing the win ter of 25, we es tab lished 2 trap ping grids [grid 1 =.18 ha (3 6 m) and grid 2 =.25 ha (5 5 m)] ap - prox i mately 15 m apart in ar eas where degus were vi su - ally abun dant. Since the home ranges of degus at our study site are typ i cally less than.5 ha (Hayes et al. 27), we as - sumed no ex change of in di vid u als be tween grids. This was fur ther con firmed by our trap ping data. Trap ping and bur row sys tem es tab lish ment We con ducted den sity-trap ping ses sions on grids 1 and 2 fol lowed by bur row trap ping in the gen eral area of the 2 trap ping grids. We ini tially con ducted 5 con sec u tive days of grid trap ping dur ing the aus tral win ters (June) of 25 and 26 us ing only large live traps (3 1 9.5 cm). We set traps baited with dry oats at fixed sta tions spaced at 5 m in ter vals [91 (13 7) traps on grid 1 and 121 (11 11) traps on grid 2] in the morn ing prior to an i mal emer gence. Af ter 2 hrs, we de ter mined the lo ca tion of all cap tures and closed the traps. We re corded the sex, body mass, and re - pro duc tive sta tus of all an i mals. An i mals were given unique iden ti fi ca tion by toe-clips and re leased at their point of cap - ture. We also re corded re cap tured an i mals. Degus are microhabitat spe cial ists, us ing het er o ge - neously dis trib uted bur row sys tems (Ebensperger et al. 29). There fore, our trap ping meth od ol ogy in volved tar get - ing bur row sys tems. In or der to de ter mine these bur row sys tems, we fit ted some adult fe males (n = 19 in 25, n = 27 in 26) with ra dio-trans mit ters (RI-2D, Holohil Sys - tems Lim ited, On tario, Can ada and SOM-219A, Wild life Ma te ri als In cor po rated, Murphysboro, IL) dur ing grid trap - ping. Us ing a handheld, 3-el e ment Yagi an tenna and an LA 12-Q re ceiver (AVM In stru ment Co., CA) or FM-1 re - ceiver (Ad vanced Te lem e try Sys tems, MN), we tracked an i - mals once per night ca 1 h af ter sun set to de ter mine which bur row sys tems they used dur ing the aus tral win ter (mid June Sep tem ber). We marked ac tive bur row sys tems a group of bur row open ings span ning 2 3 me ters in di am e ter (Hayes et al. 27) with wooden stakes la beled with a unique num ber. Sub se quent to grid trap ping and night time te lem e try, we trapped known ac tive bur row sys tems dur ing the aus - tral win ter to spring tran si tion (Sep tem ber No vem ber) of both years. We placed 1 12 traps in lo ca tions free of veg e - ta tive im ped i ment at each ac tive bur row sys tem. We de - ployed 4 trap types to de ter mine their ef fec tive ness in cap tur ing in di vid u als of 6 de mo graphic cat e go ries of degus: adult males, adult fe males, ju ve nile males, ju ve nile fe - males, male pups, and fe male pups. We trapped 34 bur row

314 J. R. Burger et al. sys tems in 25 and 68 in 26. Bur row trap ping ses sions cor re sponded with lac ta tion, which en abled us to an a lyze the trappability of all age classes: pups (25 69.9 g) ju ve - niles (7 129.9 g fe males, 7 139.9 g males) and adults (> 13 g fe males, > 14 g males) (Ebensperger and Hurtado 25). Two types of traps were solid-walled: the me dium live trap (23 9 8 cm) and the large live trap (3 1 9.5 cm). The lo cally pro duced mesh -like trap (herein re ferred to as mesh ) (3 11.5 9.5 cm) and the mesh Tom a hawk No. 21 (herein re ferred to as Tom a hawk) live trap (41 14 14 cm) al lowed for ex ter nal vis i bil ity from in side the trap. Tom a - hawks were en tirely con structed of wire mesh and were the larg est of our traps. The mesh traps ex hib ited wire mesh on the sides and rear, with solid metal tops and bases. The 3 traps had solid metal bases, al low ing most of the bait to be re tained in side the trap. Tom a hawk traps were the only traps with wire bases, per mit ting bait to fall through the trap. The trea dle mech a nism was more likely to be come im peded by oats in large and me dium traps dur ing con sec u tive trap ping days. We cleaned the oats from these traps as sid u ously be fore set ting them dur ing each trap ping event. Dur ing both years of the study, we used a pro por tional com bi na tion of trap types at each bur row sys tem for each trap ping event. We set an un equal num ber of traps, per trap type, per bur row sys tem, due to the num ber of traps per trap type avail able. How ever, the ra tio of traps per trap type re mained con stant at each bur row sys tem dur ing each trap ping pe riod. In or der to cor rect for this sam pling bias we di vided the num ber of trap days for each trap type by the num ber of trap days of the trap type with the low est num ber of trap days. We then di vided the ab so lute cap tures for ev ery trap by the for mer ra tio to ob tain rel a tive cap - tures for each year. In 26, we in creased our trap ping in ten sity more than two fold to in cor po rate more bur row sys tems but re mained within the same gen eral area of the grids. We set traps baited with dry oats in the morn ing prior to an i mal emer - gence at bur row open ings, along run ways, or nearby dustbathing sites. At ca 1.5 hrs af ter degu emer gence from bur rows, we de ter mined the iden tity and lo ca tion of all cap - tures and closed the traps un til the next trap ping event. We kept an i mals in the shade dur ing pro cess ing. As with grid trap ping, we de ter mined the sex, body mass, and re pro - duc tive sta tus of all cap tures. We re corded re cap tures and gave each new an i mal a unique iden ti fi ca tion us ing toeclips. Af ter pro cess ing, we re leased the an i mals at the same bur row sys tem of their cap ture. Our field pro ce dures were ap proved by The Uni ver sity of Lou i si ana at Mon roe In sti - tu tional An i mal Care and Use Com mit tee and are in ac - cor dance with Chil ean law (per mit no. 1-58.25-2711 by Servicio Agrícola y Ganadero). Sta tis ti cal anal y sis We did not trap degus us ing a stan dard grid de sign, there fore we cal cu lated the pop u la tion size us ing a close cap ture with het er o ge ne ity model (Cooch and White 28) for the ar eas around grids 1 and 2 for both years. This al go - rithm con sid ers the het er o ge ne ity of in di vid ual re cap ture prob a bil ity ex plic itly. When in di vid u als vary in their cap - ture prob a bil i ties, the most catchable an i mals are likely to be caught first and more of ten. This leads to cap ture prob a - bil ity be ing over es ti mated and abun dance be ing un der es ti - mated. We used four con sec u tive days of bur row trap ping dur ing the aus tral win ters of 25 and 26 to de ter mine an i mal pop u la tion size. Dur ing the course of this study, we never re corded a sin gle an i mal from the area around grid 1 to use bur row sys tems around grid 2 or vice versa. There - fore, we con ducted 2 den sity es ti mates per year us ing cap - ture data from bur row sys tems in the ar eas around grids 1 and 2. We per formed these anal y ses us ing the MARK soft - ware 5.1 (White and Burnham 1999). We used sta tis ti cal log-lin ear mod els for mul ti di men - sional con tin gency ta bles (Zar 1996) to test for dif fer ences in fre quen cies of oc cur rence of in di vid ual degus by trap type (four trap types), by sex (males vs fe males), by age (pups vs ju ve niles vs adults), and by year (25 vs 26), as main fac tors (An thony et al. 25). We ter mi nated the use of me - dium traps in 26 due to poor trap suc cess in 25 [8.77 cap tures/trap day (n = 1,312 trap days) com - pared to 22.45 cap tures/trap day (n = 4,231 trap days) with the other 3 trap types]. Since we did not use the me dium trap in 26, we omit ted this trap from the anal y - sis in or der to test for interannual dif fer ences in trap ef fi - cacy. We used stan dard ized re sid u als from the log-lin ear anal y ses to rep re sent the de gree of de vi ance from the null model (see An thony et al. 25 for a sim i lar ap proach). We ver i fied the sta tis ti cal sig nif i cance within lev els of a fac tor by ex am in ing the com po nents of max i mum like li hood, com - par ing these val ues with the crit i cal level of sig nif i cance for 1 de gree of free dom (Flaquer et al. 27). We as sumed sta - tis ti cally sig nif i cant dif fer ences at p <.5. Re sults The close cap ture with het er o ge ne ity model in di cated that the size of our degu populations com prised of 37.34 ± 1.61 (95% CI = 36.21 44.57) and 37.14 ± 2.75 (95% CI = 36.7 53.25) adults in win ter 25 around grids 1 and 2, re spec tively. In 26, adult pop u la tion size around grid 1 in - creased to 87.25 ± 8.93 (95% CI = 78.41 119.2) and de creased to 22.32 ± 3.96 (95% CI = 19.52 39.98) in the area around grid 2. Night time te lem e try iden ti fied 21 and 51 ac tive bur row sys tems suit able for bur row trap ping in 25 and 26, re spec tively. Across years and trap types, we cap tured 2672 in di vid u als in 17 79 trap days (15.1% trap ping suc cess) dur ing bur - row trap ping. In 25, we trapped 169 in di vid u als in 5543 trap days (19.28% trap suc cess). More than half of these were adults (55%), fol lowed by pups (24%), and ju ve niles (21%) (Fig. 1 and 2a), and 6% of in di vid u als were

Trapping population structure of Octodon degus 315 Adjusted captures of Octodon degus 14 12 1 8 6 4 2 22 2 18 16 14 12 1 8 6 4 2 25 26 Pup Juv Ad Pup Juv Ad Pup Juv Ad Fig. 1. Mean num ber of cap tures (± SD) of Octodon degus by trap type, age, and year of sam pling. Cap tures were ad justed to con trol for dif fer ences in sam pling ef fort within years be tween trap types. Ad adult, Juv ju ve nile. fe males (Fig. 2b). Dur ing 26, we cap tured 163 an i mals in 12 166 trap days (13.26% trap suc cess), with a sim i lar pro por tion of adults (54%), but with higher fre quency of ju ve niles (33%), and lower pro por tion of pups (15%) (Fig. 1, Fig. 2a). In 26, 53% of in di vid u als cap tured were fe males (Fig. 2b). Four trap mor tal i ties oc curred (all in 25) in 2682 to tal cap tures (.15%). These in volved 1 fe male and 1 male pup in me dium traps and 1 male pup and 1 male adult in mesh traps. The sta tis ti cal log-lin ear model for mul ti di - men sional con tin gency ta bles al lowed us to test the ef fi cacy of the dif fer ent trap types in catch - ing in di vid ual degus de pend ing on sex, age, and year of sam pling (Ta ble 1). Au to matic se lec tion of the best model via back wards elim i na tion yielded a model in volv ing four, two-way in ter ac - tions (G 2 2 = 24.1, p =.23). There fore, we re - jected the null model test ing the in de pend ence of the fac tors an a lyzed. We de tected one in ter ac - tive ef fect as so ci ated with trap type (Ta ble 1), con firm ing that the use of dif fer ent trap mod els af fected the de mo graphic es ti mates of degu pop - u la tions. A sig nif i cant ef fect of trap type (G 2 2 = 315.7, p <.1) (Ta ble 1) showed that trap types dif - fered in their ef fi cacy in catch ing in di vid u als. Tom a hawk traps cap tured ap prox i mately half of the in di vid u als dur ing both years (48% in 25 and 56% in 26), and hence, proved to be the most ef fi cient method for catch ing degus, fol - lowed by mesh traps (26% and 27%, re - spec tively), and large traps (13% and 18%, re spec tively). Me dium traps showed the low est ef fi cacy (12% in 25), but this was sim i lar to large traps (Fig. 2c). Tom a hawks yielded two times more cap - tures than mesh s, and three or four times more cap tures than large and me dium

316 J. R. Burger et al. Frequency occurrence (%) Frequency occurrence (%) Frequency occurrence (%) 1 8 6 4 2 1 8 6 4 2 1 8 6 4 2 (a) (b) (c) Medium Medium 25 26 Medium Pup Juvenile Adult Male Female 25 26 Fig. 2. Fre quency of cap tures of Octodon degus by age (a), sex (b), and trap type (c), dur ing the aus tral win ter to spring tran si - tion (Sep tem ber No vem ber) 25 and 26.

Trapping population structure of Octodon degus 317 Ta ble 1. Re sults of the sta tis ti cal log-lin ear model for mul ti - di men sional con tin gency ta bles test ing for dif fer ences in fre quen cies of oc cur rence of in di vid ual Octodon degus by trap type, sex, age, and year, as main fac tors, and their interactions. Factor Degrees of freedom Trap 2 315.73 <.1 Sex 1 15.66 <.1 Age 2 245.42 <.1 Year 1 285.11 <.1 Trap sex 2.2.99 Trap age 4 72.84 <.1 Trap year 2 2.46.29 Sex age 2 41.15 <.1 Sex year 1 4.18.4 Age year 2 22.63 <.1 Trap sex age 4 1.72.78 Trap sex year 2 2.23.32 Trap age year 4 7.41.11 Sex age year 2 6.46.3 Trap sex age year 4 1.1.89 2 p s a pat tern that was sim i lar be tween years (in ter ac tion trap type year: G 2 2 = 2.4, p =.29). Through out our study, wire mesh traps (Tom a hawks and mesh ) cap tured more than the 7% of in di vid ual degus (Fig. 2c). We found all trap types to be equally ef fi cient in sam pling sexes. Fur ther more, we found no sig nif i cant sex ra tio bi ases be tween traps (in ter - ac tion trap type sex: G 2 2 =.16, p =.99), de - spite the fact that more fe males were sam pled as a whole (fac tor sex: G 2 1 = 15.6, p =.7) (Fig. 2b). Oth er wise, the dif fer ent trap types showed het er o ge ne ity in sam pling the three age classes of degus (in ter ac tion trap type age: G 2 4 = 72.8, p <.1) (Fig. 2a). Adults were the more fre - quently trapped age class dur ing both years (fac- tor age: G 2 1 = 245.4, p <.1), but age struc ture of the pop u la tion changed be tween years (in ter ac tion age year: G 2 2 = 22.6, p <.1). and me dium traps pro - vided a sim i lar pic ture of the age struc ture of the pop u la tion dur ing 25. Half of the in di vid u als Standardized residuals 4 3 2 1-1 -2-3 Adult Juvenile Pup -4 Medium 25 26 Fig. 3. Stan dard ized re sid u als af ter log-lin ear mod els test ing the in ter ac tion be tween trap type and age class dur ing 25 (G 2 6 = 6.7, p <.1) and 26 (G 2 4 = 42.6, p <.1). Re sid ual de vi a tions higher than ± 1.96 are sig nif i cantly dif fer ent from zero.

318 J. R. Burger et al. cap tured were pups; adults rep re sented about 3% of in di vid u als. This pat tern was op po site to that ob served by Tom a hawk traps, which sam - pled 65% of adults and 15% of pups. s showed a sim i lar pat tern to Tom a - hawks dur ing 25 (with more adults than pups), which was sim i lar to large traps dur ing 26. Year of sam pling did not mod ify the age struc ture of the pop u la tion de ter mined by ev ery trap model (in ter ac tion trap type age year: G 2 4 = 7.4, p =.11) (Fig. 2a). Standardized residuals of the significant interac tion be tween trap age af ter the log-lin ear mod els re vealed that Tom a hawk traps oversam pled adult degus and un der-sam pled pups and ju ve niles (Like li hood ra tio Chi-square L 2 2 = 19.4, p <.5) (Fig. 3). The other three trap types showed op po site pat terns, and un dersam pled adults while over-sam pling pups and ju ve niles. Dur ing 25, me dium and large traps showed iden ti cal pat terns (me dium : L 2 2 = 15.3, p <.5, large : L 2 2 = 17.7, p <.5) (Fig. 3), whereas mesh traps showed non-sig nif i cant pat terns (L 2 2 = 1.9, p >.5) (Fig. 3). Dur ing 26 both mesh and large traps un der-sam pled adult degus and over-sam pled ju ve niles and pups. Dis cus sion In 2 years, we logged more than 17 7 trap days us ing 4 trap types dur ing bur row trap ping. Trap mor tal ity was low in our study in clud ing zero mor tal ity in 26. Tom a hawks were the most ef fi cient traps for cap tur ing in di vid u als, ac count ing for half of the cap tures made dur ing both years, fol lowed by the mesh traps. and me dium traps showed lower ef fi cacy. Our re sults agree with O Farrell et al. (1994) and Lam bert et al. (25), who sug gested that mesh traps are more ef fec tive at cap tur ing small mam mals than solid-walled traps. Dif fer - ences in ef fec tive ness may ac count for bi ased es - ti mates of den sity when us ing dif fer ent trap types. Tom a hawk traps were the larg est of our trap types and the only model en tirely made of wire mesh used in our study. Pref er ence for Tom a hawk traps may be due to the size of the trap open ing, as well as the spa cious ness and open ness in side the trap, which re flects the ex - ter nal en vi ron ment in which degus for age. Ad di - tion ally, an i mals may eas ily see food in side the traps, fur ther en tic ing them to en ter. We found dif fer ences in the three age classes sam pled by the trap types used. Tom a hawks over-sam pled adults and un der-sam pled pups and ju ve niles dur ing both years. This pat tern was con trary to that ob served by the other three trap mod els, which in gen eral, over-sam pled pups or ju ve niles, and un der-sam pled adults. None the less, rel a tive cap tures of pups were sim - i lar in the dif fer ent traps through out the study (Fig. 1), sug gest ing that dif fer ent traps can equally sam ple this age class. Thus, our ob ser va - tions do not sup port con cerns by Iriarte et al. (1989) re gard ing the abil ity of Tom a hawks in sam pling youn ger degus. We have no ev i dence to sus pect that ju ve niles or pups are able to es cape from the larger mesh Tom a hawk traps. Our study pro vides in sight into ques tions pro posed by Fulk (1976) and Jaksic et al. (1981) re gard ing the ef fec tive ness of me dium traps in cap tur ing Octodon degus. We found poor trap suc cess us ing me dium traps in 25, and our re sults pro vided fur ther ev i - dence that the size of these traps may be a lim it - ing fac tor in cap tur ing large (adult) degus. Adults were un der-sam pled by traps, sug gest ing that this age class did not en ter solid-walled traps readily. Since adult in di vid u - als for age above ground, they may be more in ter - ested in for ag ing in open hab i tat than en ter ing a solid-walled trap that re sem bles a bur row. Our trap suc cess de creased sub stan tially from 25 to 26 pos si bly in re sponse to a more than dou bling of our trap ping ef fort be tween years (5543 trap days in 25; 12 166 trap days in 26, within the same time frame and area), re sult ing in an in creased trap ping ef fort be yond a sat u ra tion point. We also in creased the num - ber of trapped bur row sys tems in 26 to in clude some with low ac tiv ity. The in creased trap ping in ten sity from 25 to 26 pro vided us lit tle ad - di tional de mo graphic data. The de crease in the degu pop u la tion around grid 2 from 25 to 26 prob a bly had lit tle ef fect on the over all de crease in trap suc cess since the adult degu pop u la tion

Trapping population structure of Octodon degus 319 around grid 1 in creased more than 2-fold. Care - ful con sid er ation of trap den sity and sam pling du ra tion (Conard et al. 28) may in crease ef fi - ciency when sam pling small mam mal pop u la - tion struc ture. Trap mor tal ity may be of spe cial con cern when study ing en dan gered or so cial spe cies where the anthropogenic in duced loss of an in di - vid ual may dis turb so cial struc ture. We had 4 trap mor tal i ties in 25 and zero in 26. None of these oc curred in the larger mesh Tom a hawk traps. Over all trap mor tal ity in our study (.15%) was far less than re ported by An thony et al. (25), which found mor tal ity rates as high as 2%. Check ing traps af ter morn ing emer gence, dur ing a peak ac tiv ity pe riod of degus (Kenagy et al. 22), al lowed a max i mum of 1.5 hrs the an i mals spent in traps ex posed to the sun and po ten tial pred a tor ha rass ment. Thus, syn chro - niz ing trap ping ef fort with an i mal ac tiv ity may min i mize if not elim i nate trap mor tal ity, yet still pro vide suf fi cient data re gard ing pop u la - tion and so cial group de mog ra phy. Our at tempt at tar get ing trap ef fort dur ing an i mal ac tiv ity min i mized time an i mals spent in traps, which ap par ently re sulted in very low mor tal ity. This may eas ily be ac com plished with di ur nal spe cies whose ac tiv ity pat terns are known. De vel op ing trap-tim ing de vices that re cord the time of cap - tures (Barry et al. 1989), and thus an i mal ac tiv - ity pe ri ods, may be a cost-ef fec tive way to achieve this goal for noc tur nal or cre pus cu lar an i mals. Pre vi ous stud ies found that choos ing the ap - pro pri ate scale (Bow man et al. 21) and trap den sity (Conard et al. 28) may be cru cial in small mam mal trap ping stud ies. Stud ies ad - dress ing so cial group and pop u la tion de mog ra - phy may re sult in mis lead ing in for ma tion if re search ers used a suboptimal trap type. Al - though we can not say for cer tain how the elim i - na tion of each trap type would af fect the cap ture suc cess of re main ing traps, the ex clu sive use of the pop u lar me dium or large trap may have pro vided se verely bi ased re sults in our study. It is likely that our trap suc cess would have de creased sub stan tially with the sole use of one of these traps, thus re sult ing in un der es ti - mates of spe cies den sity (O Farrell et al. 1994) and er ro ne ous de mo graphic in for ma tion. On the other hand, achiev ing the op ti mal num ber and type of trap may be eco nom i cally ef fi cient and ef - fec tive in sam pling small mam mal de mog ra phy in pop u la tion and sociality stud ies. In our study an i mal, Octodon degus, the sole use of Tom a - hawk traps may pro duce suf fi cient, un bi ased de - mo graphic data. Con sid er ation of the ap pro pri ate trap type is likely im por tant for sam pling de mog ra phy ac cu rately in other spe - cies as well. Ac knowl edge ments: We thank Universidad de Chile, in par - tic u lar, J. D. Gar cia, for pro vid ing fa cil i ties for field work, M. Hurtado, V. Quirici, C. León, D. Lahr, J. Childers and M. Pardue for field as sis tance, and T. Knowles and two anon y mous re view ers for help ful com ments on this manu - script. JRB, ASC, and LDH were sup ported by grants (to LDH) from the Na tional Sci ence Foun da tion EPSCoR (#55391), the Lou i si ana Board of Re gents Re search Com - pet i tive ness Pro gram (#LEQSF 27-29-RD-A-39), The Uni ver sity of Lou i si ana at Mon roe HHMI Pro gram and the ULM of fice of Ac a demic Af fairs. 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