Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. tpe Mirkin Stephen towns cornutum) small (Phrynosoma in lizards living horned Texas of release Predation tpe Mirkin ( Stephen lizards horned Texas for soil. 60). of refuge the = release a of provide (n Predation color may ranch the resources the match food early predators. not to and in from did compared habitat lizards location models 1) suitable horned each the have = like in when that (n species days environments especially town prey 9 common modified some in for human be attempts that field to suggest fewer the appear predation results significantly in attempts more Our and predation left significantly behind avian were observed controls left We ranch, 21) on marks the taxa. = than preserving predator On (n for models by foam ideal categorized reduced the were of is to on marks towns pieces that Texas due predation attempts control material subsequent south be a and and two may foam, summer 126) in late lower urethane towns = areas and were from (n these natural levels models Models in in predation constructed observed than whether We predators. test lizards/ha) densities by ranch. found to (˜50 high nearby be lizards higher a the still horned much on Texas be low for are than model can reason relatively much that used they one their in We densities However, declining that reach and pressure. species. been can predation hypothesized characteristics prey have We they of These lizards where loss These lizards/ha). towns and eyes. Texas (˜4-10 pressure. agriculture, the south predation urbanization, small from high increased some under to blood in due are cranial squirt range lizards sharp to eastern these camouflage, their suggests ability including of wild the predation, the avoid and in to body, rates ways survival the of number of a flattening have horns, cornutum) (Phrynosoma lizards horned Texas Abstract 2020 30, October 2 1 ffa n=2)wr eti h edfr9dy nec oaini al n aesme n subsequent 60). = and (n summer ranch late the to and compared early 1) on in = attempts (n location pieces predation town each control in more attempts in and significantly fewer days 126) observed significantly 9 We and = controls for (n taxa. on foam, field Models than predator urethane the models by levels predators. from the in categorized by predation models left were behind constructed whether were marks We left 21) test predation marks ranch. = to preserving nearby (n be areas lizards for a foam may natural ideal horned on of towns is Texas in than these that towns model than in Texas material observed used lizards/ha) south a We densities south two (˜50 high small in higher pressure. the some lower much for predation in were increased reason found are under reduced eyes. one to be that to are the that due still densities due lizards hypothesized range be from reach We eastern can these blood can lizards/ha). their they suggests However, (˜4-10 of they squirt wild much species. where to in prey the towns ability declining of Texas in been loss the rates have and and lizards agriculture, survival These urbanization, body, low pressure. relatively the predation their of high and flattening characteristics horns, These cranial sharp camouflage, ( lizards horned Texas Abstract: author *Corresponding 1 eateto ilg,TxsCrsinUiest,Fr ot,TX Worth, Fort University, Christian Texas Biology, of Department ea hita nvriyCleeo cec n Engineering and Univ Science Christian of Texas College University Christian Texas 1 ,Mr .Tucker R. Mary *, 1 ayTucker Mary , hyooacornutum Phrynosoma 2 1 enA Williams A. Dean , n enA Williams A. Dean and , hyooacornutum Phrynosoma 1 aeanme fwy oaodpeain including predation, avoid to ways of number a have ) 3 2 iigi ml towns small in living ) Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. uvvlrt 895% sas fe trbtdt ihpeainpesr Fi n ek 99 nrs et Endriss 1999, Henke and (Fair annual pressure low predators predation III Their of high 2003). Middendorf multitude to (Sherbrooke attributed a 1987, felids have often Sherbrooke and also to canids, is known 1975, (8.9-54%) rodents, are Parker rate birds, lizards predation survival specific and lizards, horned to and predatory Texas (Pianka squirting, snakes, response 2008). predators blood including: Sherbrooke in horns, distinct 1992, forces cranial Sherbrooke to selective coloration, and response these cryptic to of include: in attributed Many and sit- behaviors 2003). be 1986) myrmecophagus, Sherbrooke Endler, as can 1975, 1974, life Parker adaptations (Edmunds, for and morphological adaptations (Pianka environments and of arid variety behavioral in a living exhibit predators, and and-wait preferences dietary and ( characteristics lizards predation ( horned in models (Bateman Texas paradox, differences anole predation predation tested green urban measure explicitly on the to predation (2010) with consistent predation of models Irshick, area. results, amounts and observing foam Their lower Mcmillan or in environments. significantly natural only clay difficulty showed and studies, of the urban these use to between Of rates the Due 2016). to 2008). 1978, al. turned Guyer (Schlauch et have conditions and certain researchers Barret under many in diversity 2007, events, thrive and Pizaarro reptiles abundance generally that and in are show increase abundance Moreno-Rueda studies environments, even and some urban richness and however, in Species environments 2008); living 2018). urban (Mckinney al. reptiles Leighton urbanization et on 2007, (French with 2017). has conflicting (Berger correlated predation al. often areas negatively that are et those these impact Rebolo-Ifran in from the 2014, results predators on and al. environments conducted of et urban been lack Consequently, Shannon have the habitat. studies 2011, to Few suitable al. due of et species lack Muhly prey a 2010, some of al. for et because refugia urbanization areas to as sensitive urban act more of are may hyperabundance out predators in that areas pushed exist shown urban to are has in them research allow and species some can of Alternatively, prey that 2012). Subsequently, rates densities al. 2011). predation et lower al. high (Fischer and et anthropogenic release (Rodewald have on ecological E mainly food) an environments subsisting experience 2012, pet may predators domestic urban al. to and attributed et that trash been (i.e. (Fischer shown have subsidies environments predation 2006). have E urban of in 2012, mammals al. rates rates et al. predation and lower leading (Shochat et paradoxically well birds environments, (Fischer areas but not paradox urban urban natural mesopredators, is predation in in on urban communities differently found largely an urban act is termed done structuring may what have in predation to authors urban plays what similar that predation environments; to indicates be others that while however, urban to adapt role evidence, suggested of to Wilcove The Increasing been hallmark able 1997, has a species 2012). but Krausman some is with and understood, al. structure counterparts (Czech et community natural worldwide (Fischer their Altered biodiversity from decline greatly of 2008). differ loss McKinney can and the communities 1998, structure in the factor al. affects major et predation how a determine is important to Urbanization an be development, will human communities. by future, urban altered the of increasingly predators in down assemblages being keystone and Top habitats certain now natural densities ecological of question, With for 2006). Removal influential conservation responsible 2005). 2001). most part al. Marzluf al. in the et et 1998, be (Fortin of (Newton to Shochat thought levels one 1998, are ( trophic wolves and lower be like documented at al. to well predation species are et thought to prey Nevertheless, prey Sih is of their difficult species. on 1998, and inherently predators prey (Lima biologists of it of structure effects to making sizes community interest population nature, governing the of in processes on been observed have historically rarely predators has that are effect prey the and understand predators. predators from and between lizards match habitat horned Interactions suitable not like have did species that models prey environments some the modified for human when INTRODUCTION refuge that especially a suggest common provide results may be Our resources to soil. food appear the attempts of color predation the avian ranch, the On ai lupus Canis hyooacornutum Phrynosoma nYlosoeNtoa ak a vnipc niecmuiisadlandscapes and communities entire impact even can Park, National Yellowstone in ) r ihyseilzdlzrswt nqemorphological unique with lizards specialized highly are ) 2 nlscarolinensis Anolis ¨ otv ¨ otv se l 08.Studies 2018). al. et ¨ os se l 08.Lower 2018). al. et ¨ os nteurban the in ) Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. h imtCut ac oae ihnteSuhTxsPan crgo sapoiaey3 mNorth km 32 approximately is ecoregion Plains Texas South the within located ranch County ( Dimmit roadrunners The greater sightings), 3 = (n ) plots, study sayi our catenifer to adjacent and in lindheimeri observations ( predator cats ( of (2013-2018), sightings raccoons seasons weekly) or for field (daily periods six frequent linear active include of walk during typically course we 1600-2000, Surveys survey, the and site. 2005). each 0800-1200 Over the al. between of During et area conducted mid-August. entire are (Moeller and the and lizards search May hours, horned we of 2.5 Texas until to end apart, minutes meters the 20 2 between last spaced times people, 10 2-5 with - transects 8 surveyed is ( site hackberry of sugar consisted and ), sites ( all grass Bermuda at such grass, native types Vegetation plots plots) windmill habitat study areas. study (tumble suburban The residential 4 of grasses 2018). variety – and mallow, a Alenius (3 false parks, represent 2015, lobed Kenedy and lamb’s-quarters, Ackel lots, ha, in (especially 2014, vacant 1.22 herbs (Wall plots yards, to native 2013 study 0.054 school since 15-17 from alleyways, plots) range lizards, censused as study horned shape, have Texas 13 in We having urban - irregular for studies. as are known (12 ongoing people) served City 3,337 of Texas Karnes – County, sites and (3,299 Karnes the towns habitat. small natural in are a two City as and are served Karnes Texas Kenedy County, and Dimmit and City in Kenedy ranch Karnes hunting 1,200-ha locations: private a three whereas in environments, models placed We Sites Field models lizard horned Texas town. adult attempts? in models and METHODS predation juvenile, foam lower of hatchling, on be do number attempts a 2) would predation their sample and do would in rates areas?; 1) landscape, differ predation natural asked: the we and that on Specifically, urban placed hypothesis lizards. between when horned them differ the that Texas placed of models, test and predators accurate to lizards of morphologically horned variety habitat create Texas to natural adult was exploitation and a goal the juvenile, Our and hatchling, and of towns lizards models small of foam densities in created we high study both this town facilitated In in has lower are which this rates items. lizards, that predation prey horned and that Texas small hypothesized areas, for of we natural therefore, time more predation; ( handling and of in ants increase risk than (Whitford may higher smaller at lizards/ha) items eat them prey (3-10 predominately put smaller would areas towns on these Foraging natural 2018). in more (Alenius Lizards in 1993). research densities ( al. termites Past reported 52.32 and Texas. et of the southern Whiting densities than in towns, average 1979, curio Texas City at higher Bryant small and occur Karnes much some towns pet and in is these the Kenedy found in which for of still lizards over-collecting towns are that lizards the and shown horned has in ants, ( Texas ants fire occurring 2003). harvester Henke with populations of 1994, including competition loss al. the and et a lizards, 1994, insecticides (Donaldson horned to al. trades of of attributed et ( young use ants are (Donaldson and fire Texas widespread lizards eggs red of common to the invasive state once conversion, on prey the habitat these eastern can and in in in which urbanization Declines species especially including: 2012). threatened factors declined, TCAP a of have – now variety southwest, Plan are American Action and Conservation the range Texas historic of their vertebrate of iconic areas an lizards, horned Texas 2020). al. et Miller 2007, al. rco lotor Procyon n=4sgtns,cahhp ( coachwhips sightings), 4 = (n ) : eursrtre cinereus Tenuirostritermes n=1sgtn) rdtr id r lorrl en e-aldhws( hawks red-tailed seen, rarely also are birds Predatory sighting). 1 = (n ) avsrmcoromandelianum Malvastrum yoo dactylon Cynodon etslaevigata Celtis .W aefudvr e nks nldn ea a nks( snakes rat Texas including snakes, few very found have We ). hoi verticillata Chloris r h otcmo re ntesuypos(al21) Each 2014). (Wall plots study the on trees common most the are ) hnpdu album Chenopodium ahrta hi omnypeerdpe flrehretrants harvester large of prey preferred commonly their than rather ) eccy californianus Geococcyx .Hnymsut ( mesquite Honey ). atcpi flagellum Masticophis n rpclamaranth, tropical and ; ei catus Felis lisbitegrass bristle plains ; 3 rspsglandulosa Prosopis ,dg ( dogs ), tage daisy, straggler ; n=2sightings). 2 = (n ) n=1sgtn) ulsae ( snakes bull sighting), 1 = (n ) ai uu familiaris lupus Canis ± eai vulpiseta Setaria 12S iad/a(ce 2015), (Ackel lizards/ha SE 11.2 mrnhspolygonoides Amaranthus aytcru vialis Calyptocarpus ,aau ( anacua ), Pogonomyrmex oeossinvicta Solenopsis ue jamaicensis Buteo lpeobsoleta Elaphe n h non- the and ) hei anacua Ehretia ,adnorthern and ), Pheidole Pituophis p)due spp) three- ; and ) spp.) ), Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. da iefrmntrn aua rdto nteelzrs u dhcosrain fpeaoso the on predators of including: observations lizards an horned hoc Texas it of ad predators making Our potential lizards of lizards. number horned a these ( Texas included hawk on 2018, Harris for of predation prey summer the natural and during monitoring ranch predators for both ( site of mesquite ideal communities honey by natural dominated maintains is habitat habitat The area). management glandulosa (wildlife WMA Chapparal the of ( h ac nDmi ony hseprmna einalwdu ots o n ieecsi predation in differences any for on test found were to that us lizards allowed and design soils the experimental red During This the time predation. County. environment. color-match two urban possibly Dimmit to these the repainted and in in used were lizards weather ranch We and models in the substrate days. summer, the differences nine late color-match for to the for painted account During setting were to models ranch environment round summer) urban natural summer, the late early the to in and placed to 4-August-2018 were (early relocated and models periods subsequently period, 29-June-2018 time were to each then of 9-June-2018 metallic days and intervals: the nine cover first time the to two over During 21-August-2018. painted during head scat. experiments nail ranch their conducted the sites and the We with 7 lizards on nails for The inch placement that surveys types. 2 model by areas using habitat determined of substrate in both as the surface. sites playgrounds town in activity to 7 in school pieces lizard secured The horned control were sites and Texas Models surveys. 21 alley-ways, 7 known and previous lots, across with models vacant from areas controls 126 yards, included determined 3 of included lizards and total City horned adults) a Karnes contained juveniles, for and (hatchlings, ranch Kenedy class the in size on the in each sites them 7 of placing and models to prior 6 days placed 7 We as, of period well a them as for covered lizards, dissipate Experiment and to Predation local fumes were outside resemble paint lizards models allow accurately where dried to would soil substrates placed of field. that we the layer The painting, models of loose substrates. After a photos the surrounding with substrates. take for the each the to of colors from color-match used taken select lizards background was photographs to horned iPhone from Texas found for as adult previously well (X-Rite) the multiple as app in of the specific placed, found Studio The photographs where soils. were PANTONE population using City red a models determined by Karnes from largely where were Texas characterized and lizards location models is of colored Kenedy substrate the variations red the of of color where and environments County colors color, distinct Dimmit urban in in two the setting gray match ranch predominately from natural to but lizards paint varied gray acrylic is 1); substrate using and (Fig. pieces paint material lizards control acrylic foam horned and the with to models painted attracted painted and being We predators models 1). for (Fig. lizard control from paint the producing 3 to constructed the casting were models of iT! to controls lizard from Foam capable or The the over casting. hours. like molds left 2 exactly per create for foam treated models cure to otherwise 3D urethane multiple to and models to allowed of and SVL), and printed used pieces study molds mm was 3D round the this (50 into that these juvenile in poured SVL). used SVL), (.obj) was used length, We mm (Smooth-On) file classes snout-to-vent (23 models. size object hatchling mm size different an lizards: (84 SVL) the create horned Texas mm lizard to model (84 scanned horned 29NV of adult Texas was classes Max replica adult size Mold lizard three an using horned print of pewter molds replica original constructed pewter The 38 We a effective of proven 2012). and has excess Forstner that (Smooth-On) in material and a temperatures (Farallo foam, summertime predation urethane Texas using controls withstanding and at models lizard horned constructed We Construction Model ( ( rattlesnake ac sparverius Falco yxrufus Lynx and ) ,cyt ( coyote ), rtlsatrox Crotalus aaue unicinctus Parabuteo ,getrrarnes n ogredsrks( shrikes loggerhead and roadrunners, greater ), Acacia ai latrans Canis : : hrsrbcmuiistpclo ot ea huln.Ti eaieywild relatively This shrubland. 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Data may be preliminary. itre ynnpeaos(..ho ak)(=)o etmsig(=)wr nyo h ac n were and ranch the on analysis. only the were from (n=6) excluded missing and went events or predation (n=2) as marks) counted 1.13x10 hoof not = (i.e. (P than non-predators ( ranch by periods models the 1.05x10 disturbed sampling adult on = attacked two or (P were juvenile the 31 ranch than over and 60) the town = on 1.12x10 in (N attacked = attacked than ranch (P were models 1) the 29 both the = at over and on (N controls models higher towns the on were the on events events in 30) occurred predation Predation = events ranch. more (N predation ( the observed Four overall summer on We 1). both early models. located (Table controls, between all were 0.89) the which occurring of = on events of 12.1% (P predation all representing were 31) of periods, periods = that number sampling sampling (N total models both summer the and 3). over late in (Fig. recorded (ranch and 0.019), difference 0.00013) color-matched were = no = events not P found P were predation We 18, that 61 =18, = those of df df than total that 2.59, 4.86, scores A those = = COI (t than higher t red), scores had ranch red, COI ranch vs higher urban the gray had on vs environment color-match gray urban to (urban the painted in color-matched color-match not to were painted were that models. Models red exact ranch Fisher’s and checked gray using and ranch analyzed RESULTS and models then models between were red color-matching data urban in and 0) differences gray of determine urban values between to assumptions expected differences t-tests the (i.e. for Where two-sample met. met used were We not natu- test and that were tests. urban of in tests assumptions placed the Chi-square models when of of tests rates Chi-square predation using in environments differences ral compared we experiment predation using percent our For (COI) the index indicating overlapping scores, color COI models. Analysis: calculated a 40 We Statistical create for v. 2015). to model, 3D Francini and model. used Inspector and Color substrate the then (Samia between the to were Rstudio color-match using (˜1cm) values in photo adjacent Function color cropped substrate COI These each from of the from ImageJ. cropped crop compare obtained was for to sized were coloration plugin possible values exact lizard’s color it 2.3, an model blue) making with the green, colors, compared that of (red, true and profiles portion RGB their ImageJ a camera (DNG) to using calibration, ColorChecker calibrated negative photo After the were digital photos. (X-Rite the and that all created 1.1.2 images standard across we create version color values to software Classic, color this used with Lightroom Using photo- be frame. Photo Adobe we then the Passport could for substrates in ColorChecker plugin surrounding USA) a software 49512, their with calibration MI color-match field Rapids, to the Grand painted in Inc. successfully model were each 2). models graphed but (Fig. removed whether pieces limbs multiple test had in To that found were models Unknown- that or models shaped and model; half-moon Color-Matching: marks marks the distinct bite peck of left Rodents- or that sides decapitated; bite bites lower clear Other- been and no model; had upper the that Bittner, both on models on and left and impressions imprints (1993) categories: marks teeth 4 Brodie, chisel peck into distinguishable categorized as shaped were predation with criterion “U” models of the or evidence similar on had “V” used marks that Birds-obvious Predation We 16-300mm models events. a Tamron removed. predation and a discovering categorizing period and app when with day upon (2003) photographed ArcGIS camera 9 and the were SLR for during placement digital disturbance Collector days d3300 initial three or the Nikon every upon a models using checked with models We model disturbance lens. photographed each or also event of We placed. predation coordinates were accuracy. potential they the 1-m which recorded with upon we (ESRI) substrate deployment, the and initial models Upon between color-matching background to due rates χ 2 χ 2 .4 f=1 .5 n nternh( ranch the on and 0.05) = P 1, = df 3.74, = .8 f=2 .2)(al ) oesta a vdneo being of evidence had that Models 1). (Table 0.029) = P 2, = df 7.08, = -18 Tbe1.Drn h al umr oe a takdi town in attacked was model 1 summer, early the During 1). (Table ) -8 Tbe1.Drn h aesme,n oeswere models no summer, late the During 1). (Table ) 5 -10 .Hthigmdl eeatce less attacked were models Hatchling ). χ 2 .4 f=1 0.04). = P 1, = df 3.24, = Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. ofidtercytclyclrdpe.Orrslssoe htwe oeswr ane obackground to less painted significantly acuity were were visual events models their predation when use avian that may cryptic), showed lizards more horned were results of they Our predators (i.e. avian prey. surroundings and colored their 1976) cryptically color-match lizards al. their horned et flat-tailed find (Fox in to visual lengths to highly horn similar are cranial lizards is Birds increased horned which of of 1990). numbers evolution (Sherbrooke lizards, large the ground horned consume for the Texas shrikes “centrifugal-slam” on responsible like ( to limbs. it predators roadrunner be threat avian smashes of even greater that and significant may loss show typical tail and a that the a or studies are caused limb other of several predators any what in result by found avian findings unknown lizard the also suggest is a We be results grabs it also bird These although comm.). the areas may limbs, which in limbs pers. missing in lizards missing attack horned Wildlife, with Texas with ranch and off models limbs Parks the the Texas The on chew Rains, lizards to (Nathan shown or adult been bite common live have predation definitive are and recorded no 2007) both but of al. where removed ( 48.3% et limbs mice Endriss represented had 1986, models grasshopper which that (Munger of Northern unknown, models venter included as marks. and events scored peck dorsum predation were unknown the chisel categories These on obvious ( those events. impressions included lizards Bite into bite Bites predatory events. falling half-moon attack models. of marks. predation not and in indicative to peck of discern incisors possibly tend 30% shaped to rodent were birds for easy “U” by that accounted relatively because or behind predators the also predation “V” Avian were left is avian conspicuous and marks 1973). concern as 21.7% of (Smith for categorized common presence prey accounted also their the a marks 1993, behind of were by 2012), left head (Brodie decapitated discern Marks Forstner the studies been to towards categorized. and other had easy accurately Farallo are were that in models 2010, birds Models predation on by behind measuring al. models left at marks (Alenius on et predation effective items Vignieri which proven prey with 2010, has confidence small Irshick consuming models and to of by McMilan shift lizards town use dietary horned in Texas the their of pressure Although density and predation high 2015) result relaxed the consistent (Ackel to and a factor City are 2012) contributing are dogs Karnes 2018) a communities and al. and be cats predator rarely may et pet Kenedy Altered birds are and Fischer in like Feral ranch. snakes predators 2004, ranch. the and of the Gehrt on types prey at and occur certain daily of is not seen (Prange birds were there do urbanization but instance, abundance but densities, towns of For town the the predator in in both ranch. measure plots common in the study explicitly predation differs also our to not City lower near compared Karnes or did found when in and we have seen predators Kenedy Although that than in of community lower studies diversity 2012). predator be other and the would with al. that town consistent et evidence in are (Fischer anecdotal lizards areas and horned Texas environment, urban on ranch in predation natural that the hypothesis in our supported results 5). Our (Fig. pads cactus on bites DISCUSSION tortoise the to shape and ranch the size on of soils the pads color-match and to ( painted pear were prickly shape round ( dried half-moon second tortoises resembled conspicuous the result during had a controls August as Our in and attacked 5). were (Fig. that marks controls bite remaining three the whereas on marks, soils background red ( to rodents:( the unknown painted ranch color-match or were to the models painted on when not 5) categories = were remaining (N rodent, models summer lizard birds, late when ( during category: 13), than color-match higher = predation significantly (N by was summer ranch events early the predation the of in birds number by the ( in unknown and difference other, significant a found We hyooamcallii Phrynosoma ohrsberlandieri Gopherus Opuntia χ χ 2 2 .,d ,P=00)(i.4.W i o e n ieec nclrmthn nthe in color-matching in difference any see not did We 4). (Fig. 0.05) = P 1, = df 3.8, = .,d ,P=07)(i ) h taki ueo oto ic a itntpeck distinct had piece control a on June in attack The 4). (Fig 0.78) = P 1, = df 0.1, = χ p.o h imtCut ac n h iemrso h otoswr iia in similar were controls the on marks bite the and ranch County Dimmit the on spp. Mne 96 on ta.2004). al. et Young 1986, (Munger ) 2 32,d ,P=2.86x10 = P 3, = df 33.24, = Fg ) efeunl nonee ea otie aigbt h fruits the both eating tortoises Texas encountered frequently We 5). (Fig. ) ncoy leucogaster Onychomys χ Opuntia 2 29,d ,P=00) te ( other 0.09), = P 1, = df , =2.9 rtpyu reticulatus. Crotophytus 6 -7 p. asadfut;wihmyhv trce Texas attracted have may which fruits; and pads spp.) Fg ) eas on httenme fattacks of number the that found also We 4). (Fig. ) r nw rdtr fhre lizards horned of predators known are ) .Alohrdmg omodels to damage other All ). χ 2 15,d ,P=0.23) = P 1, = df , =1.5 Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. ygat rmteAdesIsiueo ahmtc cec dcto tTU n rnsfo the from grants and TCU, Karnes at the Education in making Science lizards and & horned Fund. ideas Mathematics Texas Activities of project Creative of funded Institute with and was studies Research Andrews project helping our This TCU the for feedback. of and from Rhoads suggestions invaluable grants support Dusty gave Watson by the and and Charles at and help Alenius Hale hospitality Amanda complete Rachel for and models. could lodging thank Phelps we for that also Wade Hunt, so We Truett and home County. his and Kenedy and Sue˜nos Maggie Ranch in Los Breckenridge, to Heatherjo 505 access thank the us We in giving project. available for this Griffin be Alan also will Mr. thank and We figures and table, text, the within ACKNOWLEDGEMENTS found be can repository. paper data this DRYAD in used data All experience. the might increase help STATEMENT release site may ACCESSIBILITY the reintroduction DATA sites lizards. to the reintroduction reintroduced prior potential pressure for at established predation survival community are by relative of predator environments they future probability the the the natural where and gauge in predation areas in tool to of in when conservation lizards Knowledge lizards a and consideration horned as horned areas conservation Texas used Texas reintroduction important be of that potential an also may in suggest be food Models them further should and placing reintroduction. Feldman and results potential and vegetation pressure, for (Keehn Our predation predators areas suitable from targeting of have 2020). species conservation levels environments prey inform high some al. modified for help experience refuge may et human a environments like Law provide these urban populations, cases 2018, If in some lizard in persist horned may to some they populations. how able resources other Understanding are study City, This for studies. environments, Karnes natural 2013). future efforts and more for al. and Kenedy et foundation urban Wolf in in 2007, a those lizards as al. horned et serve Texas Endriss of may one 1994, predation and as comparing al. suggested data et often first (Donaldson is declines the urbanization those presents and of prey. range drivers potential historic main their as the may of throughout the them declined result constructing Conversely recognized have in have This lizards difficulty predation. horned not and Texas of size simply models. small types therefore their adult may certain to predators or due to realistic, and juvenile vulnerable least them less the the morphologically are we than were encourage lizards future, models pads. attempts hatchling to hatchling cactus the predation cues stationary for In fewer visual them that had mistaking no models. indicate tortoises offer models our Texas would hatchling construct by that to occurred Our pyramids) predators have used may (i.e. suggesting paint however, as shapes or such controls into events foam the made predation urethane than be the more controls (Bateman significantly that to 2017) models attacked recommend attracted Marra the of were construct limitation simply and to models Another (Loss not used Our events. material areas were particular predation known 2016). urban the these are to sample in Cats al. attracted to mammals are et house. suited predators and the well some that to birds, be be for back may not reptiles, areas models lizards may small horned studies urban residents of dead model the local bring stationary predators Nevertheless, attracted in and will efficient often models. cameras cats very the are pet video ignoring be they their detection and to since exhibited sometimes by motion that cats not walking up us by cats are told attacked set of which have We instances be of predators several rely to all filmed 2008). snakes of 1983); models and Wells because Buning models subset expect and snakes Cock not by (Ellis a (de attacked did motion prey be sample we by movement, Similarly, to sense models models to of lizards. expect cues therefore lack model not olfactory by the and did and we include motion, responses instance, which thermal, For behavioral studies on 2016). predation appropriate al. for et of (Bateman models lack visually using and against in smell, limitations lizards 1975). inherent horned Parker crypsis are and for that Pianka There adaptations hypothesis 1964, long-held Lowe defensive a and primary supports (Norris the This predators are color-match. oriented to color-matching painted background not were and models when than likely 7 Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. o,R,Lhkhe .. n etnof ..17.Flo iulacuity. visual Falcon elk 1976. influence D.H. Wolves Westendorf, and 2005. Park. S.W., J.S. National Yellowstone Lehmkuhle, Mao, in R., and Fox, cascade T., trophic Duchesne, paradox: a D.W., predation shapes Smith, the behavior M.S., Boyce, and movements: H.L., Urbanization communities. Beyer, vertebrate 2012. D., structuring Fortin, J.R. in Miller, dynamics trophic and habitat of T.P., a role Lyons, in the S.H., polymorphism Cleeton, color J.D., of maintenance Fischer, the and ( Predation lizards squamate. 2012. horned specialist M.R. Texas Forstner, of and survival V.R. and Farallo, ranges, home Movements, with 1999. cornutum pressure S.E. soma predation Henke, of reduced and population indicates W.S. urban meta-analysis Fair, an A of 2018. Demography G.L. Lovei, urbanization. 2007. and increasing W. T. R. Oklahoma. Magura, Moody, central C.B., & in Eotvos, cornutum) F., (Phrynosoma S. Fox, lizard Chicago C., horned of Texas E. University the Hellgren, Prey A., Vertebrates. Predator Lauder, Lower D. G.V. Endriss, of a and Feder Study in M.E. U.S.A. the housed In Illinois, from cats 109-134. Chicago, Approaches of Press, Pp. and behaviour predators. Perspectives the against on Defense Relationships: stimulation 1986. visual A. New J. of York, Endler, influence New The Longman, 2008. defenses. shelter. Texas D.L. rescue Wells, anti-predator the and of in of S.L. survey prospects feeding Ellis, A future and Animals: and capture in status Defense U.S.A. current prey York, The 1974. for M. 1994. specialization Edmunds, J. a Morse, as and ( lizard A.H., sensitivity horned Price, Thermal W., United Donaldson, the 1983. in endangerment T. species Buning, snakes. of causation Cock and de Distribution 1997. P.R. predators Krausman, States. avian and free-ranging by B. patterns Czech, banded snake coral of Rica. avoidance Costa Differential in 1993. E.D. III, Brodie of Science models of clay Journal areas. Polymorphic Ohio protected 2003. in T.D. predators and Bittner, of prey use of redistribution the the modelling: and ecological shields Letters human of Fear, vertebrates. kind 2007. in J. different interactions Berger, predator-prey A stream explore 2016. and to A.K. riparian organisms Wolfe, in model and reptiles clay P.A., and USA. Fleming, Georgia, amphibians P.W., western of Bateman, in responses disturbances Differential use 2008. land C. to habitats Guyer, and K., Barrett, ( lizards horned Texas University. of Christian analysis Texas horned Diet Texas 2018. of R. management conservation Alenius, for estimation population details: the ( in lizards devil The 2015. A. Ackel, REFERENCES Science mrcnZoologist American :620-623. 3: hyooacornutum Phrynosoma hyooacornutum Phrynosoma ple nmlBhvorScience Behaviour Animal Applied Evolution 7:1116-1117 277: ). ora fHerpetology of Journal lSOne PloS adcp n ra Planning Urban and Landscape 0() 62-66 103(3): 71:227-235. 47(1): 32:363-375. 23(2): () e30316. 7(1): .Mseso cec hss ea hita University. Christian Texas Thesis, Science of Masters ). nTexas. in ) 517-525. hmohssirtalis Thamnophis ea ora fScience of Journal Texas 1(-) 166-174. 113(1-3): , 8 hyooacornutum Phrynosoma 8:54-59. 180: ilgclConservation Biological Bioscience ugs atrso va predation. avian of patterns suggest Herpetologica ora fZoology of Journal 6 97-113. 46: Science Ecology .Mseso cec Thesis, Science of Masters ). 2 809-818. 62: 4:2290–300. 141: 9(26:263-265. 192(4236): 33,320-331. 63(3), , 6 1320-1330. 86: 0:251-262. 301: Phryno- Biology Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. 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Phryno- Urban Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. n ymdlsz ls n2018. in class horns. size its model got by lizard and horned the imperiled 1. How Table to 2004. threats E.D. Quantifying Brodie, 1998. and E. K.V. Young, Losos, and States. A., United horned Phillips, the Texas J., in of Dubow, species population D., Rothstein, a D.S., of Wilcove, distribution Spatial 1993. ( R.C. ralist lizard Murray, horned and the ( J.R, prey: lizards its Dixon, and M.J, predator Whiting, a of Behavior ( 1979. University. lizards M. cornutum Christian Bryant, horned Texas and Texas Thesis, W.G of Science Whitford, of genetics Masters and Texas. range south mice. Home in in towns crypsis 2014. of advantage A. selective Wall, The 2010. H.E. Hoekstra, 2153-2158. and 64(7): J.G. Texas. Larson, Editor, Austin, S.N., Overview. Vignieri, Coordinator. 2012-2016: shrikes, Plan Plan loggerhead Action Action Conservation Conservation young Texas Texas Connaly, 2012. in Wendy Department. behaviour Wildlife and prey-attack Parks prey. Texas of on study predators A multiple of . impacts 1973. Emergent S.M. Smith, 1998. D. Wooster, Evolution and & G. Ecology Englund, processes A., emerging to Sih, patterns From different 2006. ecology. with D. Hope urban taxa and mechanistic N.E, snake McIntyre, in two S.H, Faeth, to P.S, Warren, lizards E., Shochat, horned Texas Press, strategies. by California subjugation responses of and Antipredator University foraging 2008. America. W.C. North Sherbrooke, of lizards horned to Introduction lizards. Berkeley. horned 2003. on W.C. feeding Sherbrooke, roadrunners greater captive of Bulletin behavior Wilson Predatory 1990. responses ( W.C., Behavioral lizards Sherbrooke, horned 2014. in posture K.R. head Naturalist Crooks, Defensive Southwestern and 1987. L.M. W.C. Sherbrooke, Angeloni, shelter. predator A.R., human-mediated Hardy, a Kirk- with L.S., associated In: Cordes, Island. G., 25–41. Resources Long Natural Pp. Shannon, and of Forestry of Service. reptiles Department Extension and (IN): Lafayette Cooperative amphibians West the the people. and and of Wildlife ecology using editor. geographical contrast CM, patrick Urban animal-background measure 1978. to F.C. Schlauch, method affordable An 2015. images. R.B. digital Francini, and D.S. predator- Samia, decouple subsidies resource Anthropogenic 2011. D.P. relationships. Shustack, prey and L.J., Kearns, A.D, Rodewald, Behaviour 8 150-154. 38: hyooacornutum Phrynosoma rdto vnsrcre ntw n nteDmi onyrnhfrtetosmln periods sampling two the for ranch County Dimmit the on and town in recorded events Predation n avse ns( ants harvester and ) ue1162/2 30/252 29/126 1/126 Total Events Predation Ranch Events Predation June Urban Experiment Predation 113-141. nentoa ora fFuaadBooia Studies Biological and Fauna of Journal International 0() 171-174. 102(1): clgclApplications Ecological 39:350-355. 13(9): 32(4):512-515. Bioscience rnsi clg n Evolution and Ecology in Trends hyooaia)rltv ohbttadprey. and habitat to relative Phrynosomatidae) : Pogonomyrmex ora fHerpetology of Journal 8 607-615. 48: 1 936-943. 21: spp.). 10 LSOne PLoS Ecology 42(1):145-153. 1 186-191. 21: 0 686-694. 60: () e94630. 9(4): Phrynosoma hyooacornutum Phrynosoma :8-16. 2: Science ara Iguanidae). Sauria: : h otwsenNatu- Southwestern The 0(67:65-65. 304(5667): aisludovicianus Lanius ntosmall two in ) Phrynosoma rnsin Trends Evolution The The Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. iue2 ersnaieeape fpeaineet nmdl trbtdt id APcs B- (A-Pecks, birds to attributed models on events predation (F). Unknown of and (D&E), examples Other (C), Representative Rodents Decapitations), 2. Figure possible as accurate hatchlings, morphologically as models (E). be of lizards to classes to horned created size painted Texas were 3 were models actual Models used (B) Gray We (D). mimic models controls study. red to ranch. and County and this (C) City Dimmit in adults Karnes the used and and on juveniles, were Kenedy found in models lizards found lizard horned lizards Texas horned gray match Texas match of to variations painted were color (A) distinct Two 1. Figure otos04 /24/84 29/84 26/84 31/252 6/84 4/42 28/42 26/42 6/42 31 0/42 1/42 0/42 0/126 0/42 Controls Adults Total Juveniles Events Predation Ranch Hatchlings Events Predation August Urban Experiment Predation 11 Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. acigi nipratdfnemcaimaantpeaoswt ihvsa ciy eosre no observed We acuity. visual categories. high models predation with when remaining color- predators the ranch background amongst against suggesting the events 0.05), mechanism on predation (P= defense predators in cryptic) important avian difference less were by an events (i.e. is color-match predation matching background more to painted significantly not observed were We 4. Figure Models scenario. than color-matching scores each COI in higher match. models significantly to 10 have painted for environments not their scores were color-match COI that background for those to error painted standard were and that Mean 3. Figure 12 Posted on Authorea 30 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160396413.33761655/v2 — This a preprint and has not been peer reviewed. Data may be preliminary. oto icsfo aesme nAgs,21 nteDmi onyrnhwt iemrssmlrto similar marks bite with ranch County on Dimmit feeding the tortoises on Texas 2018 (D) ranch. August, A, in in summer those late from pieces (A) control 5. Figure Opuntia p.pd ihbt ak rmTxstross( tortoises Texas from marks bite with pads spp. Opuntia 13 p.wr rqetsgtn nteDmi County Dimmit the on sighting frequent a were spp. ohrsberlandieri Gopherus (B&C) )